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Contributions
of the
American Entomological Institute
07 AUG 1967
"Volume 2: Numer 1. 1067
—"
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA. - Il
The genus Culex in Thailand (Diptera: Culicidae).
by
Ralph A. Bram
CONTRIBUTIONS
of the
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Contributions -
of the
American Entomological Institute
Volume 2, Number 1, 1967
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA. - I
The genus Culex in Thailand (Diptera: Culicidae).
by
Ralph A. Bram
CONTENTS
PE EPROOGUG TION i ee es i ee ee Pw ev oe 2, ad eg Poe 1
CP Oe i nh be be ee ua a's ghee gen ya i, Saree an 7
Keys to tie Submenera in THAUANe: 6h ee ees 9
eS Pe Ae ie a ee We Pie Gla ee ek els he ee 11
Keys 10 the species im THAUAMe ys ee ee ee 12
TUS CONUS WiGGOIAR oe. he A Ce a ee we 13
HOMER TWOCOMAIG. 6 a a as ii etka a gy ae 17
DUE CRUS NRO UN ec ee ee Be A ek owe bk he 20
Keys to the species in Thaland. 2 ee i ee i ee 24
OOO ey Owe ee hele es 25
ER ee TO ee es Gere ea bee we eS 29
mune CT ONE ete EN es al a ee ees 33
Ove (0 the ByeCICs in TRAIAN ok 6 is oe ene ea ee 34
Peers OM a a Fhe ey Pee DS ak es 34
eee OG pine ee ee be ee ae as 37
De OIC OMTET oe ie i ee ek ie 39
Suorenue LOPHPOC RRA OMT Ar ea sere Peg Oe se Ba ws 42
evs to the Gnceies in Tiatand i Oe ee es 44
creme Re ee a ee ie ee a eh, Ce ew 49
Ot ee eee i ee ee ee Boe ew 20
Clee COUeee ee Sis ts A er D3
CHAUIC LS TOMY rey Ps eek a es 20
eS PA e ee a ee ek ee eas 58
WICCUOE AT COUOES 6 6 ee a a OS A 61
WIG TPG IPis TOWERS) a ee oe aS . 65
Ve CON ee a ee a 66
PUUUHOVACTS MIMEICESTON) i i a Bg ew ee ee 69
Pe APMC i a ee eas ees 72,
Momucie: Oe ee ia 75
PEO PS UO a a lala ee wee 15
CUTS TOW i kee ES ob ee es 76
Te i Wa ke See we es 79
Wer OT ON a era ee ke ea ee 81
incomptus Bram and Rattanarithikul............. 81
RRC TON SS TORN AU eg A ek a a ee 82
eukrines Bram and Rattanarithikul............... 85
fuscosiphonis Bram and Rattanarithikul........... 87
SUING, COOLIORR ee aia a ke ee ee ae eae eae 90
Hiner eT (RIC OGLON) ba a ek em ea 93
EO Te Oe a ea 96
minor form bicornutus (Theobald)............... 99
peyiont Bram and Rattanarithikul....-....2...0.%54. 100
bholeter Bram and Rattanarithikul...... ee ae: 103
spicuiosus Bram ane Ratianaritniktl. . kk ee 105
BOVE CET a SVG) Fue abies a gke la e 108
WIT ORT COTM i eee eet 110
PPOs TAO a ee ea 113
PSE R TUS OR er yk area ay ws ky ick a 113
il
Submenus COULICIOM Eo ee i Bie hk eae ee eae 117
Keys to species In THATma re bis Geile wi ee Bee le bo ea ey 120
OAC E DBP VANE be The yi ee a a RS 123
COTTE DB il igre ie maw ee Oe Gt 125
JVELUS TABOR 3 eB EN EA ORES a VG 129
NAST OPUNCTAL US POWALCB kin ae ep wo wn ei pt Oe 133
palligotlovas:TROORAI 4g SO er ay ee he 137
papuensis (Vaylor). hanes ke Ow RG 143
SCORE BE By OG ie Bee Wb eeu ic Bs 147
SOOTY CO ATOR ie i Li ale ME es RE ei 151
SOCCULOLIEOTEN Wie) BR FO a Oa iG hg cgi lial ia anata MIE 155
Lerid TW RE Ce ee 157
FRU THIQROVIN BO a LG alos Ste ar ar tanea er Mar Gene 159
oa cg hr a Oe a ay. Gama mr MIEN ci! SSD ah ROM as i aa Meas eR NICO bol aie rine to) si Pe 163
Keys to species in Tiara 7 re ee uae a eae wa 169
POTN PO ae OR ee ae ee 182
TRSCOCEP Pdi TROCHRIE i SE OW ees 184
WECR SG BATT oe EEN PR POP EG 188
PIPIENS QUIGUCTASCTIATUS Say Oe i Pee OR 192
BICC Se MEE Es OE EOP ie Pe EI a Se ee et ', 201
VISIO SAPO Oe VE Eee So ree Oe a 202
alienus Colless ...... ee SEPT So WN Mh cg OCT ag 202
CATS THOOIGER fe Fa SO ae Wa es ek 205
DOV? BEWar Ge ee we OS EO 209
HUMMUS SHOE oS OES PEW A ek Bwana 213
perplenus Leicester. oy ek es eae Gur are 221
PSECULOUISAHU TOLLESS PSS Oe A wa 222
LVELCCHIOVN VRCHUS CAGE 8 I ES ERs 225
WHET BAPE ORs eS YO 8 WEBLY OEE SS 233
Sei ens Baer Ouse ee iy Be ee We a, a ees ie ony 236
REHAMOTONS BBO PE RUS ORS ES ee 236
SUNS WIOdeTiaA i ee Be EN ew 239
LOLS: FO EEG WORT TGS, oe BO RS aa OE 247
gelidus Theooald 2303. De 6 ek Oh arg kd 247
WILTED) 6 OOO AES es 253
BUACCRTOVR INTIS BAO OOD Ni a OR AEE 257
DIM CHIOTHIECUS GIES iE ek ee A es OS ee 258
PSCUMOSEICNSTS CLO OR Bo OE i wie Wee CN PG 265
SHLCNCIS: LNOOORIG BP We ee i oe VE OBES 268
WUSIDENTIFICA TIONG 22028 OF SA Ga Ge Oe a er ae 274
C. (Mochthogenes) castrensis Edwards........... 274
C. (Lophoceraomyia) flavicornis Barraud......... 274
C. (Lophoceraomyia) fraudatrix (Theobald)........ 274
C. (Lophoceraomyia) uniformis (Theobald)......... 275
C. (Culiciomyia) viridiventer GileB i. es 275
C. (Culex) pipiens pipiens Linnaeus....... AG ie ae
, Ci Glules) Heelers Theanale sae ais ewe 275
en ee ee each a ag cet its ou peor 275
C. (Mochthogenes) khazani Edwards............. 275
C. (Lophoceraomyia) minutissimus (Theobald)...... 276
C. (alten) Cornnins SOWAGR ie a 276
lil
GC. (Cuber) mae 06 i Pn i IE 276
C. (Culex) wunivibdins Tnegoald ... a 276
©. (Cdlex) vishnws Theopale iv. ee 276
POINOW DNS oe 2 eae ak ee Oe ee ees 277
De UO OU an a ee ee Ge a 278
BP a eh eA he ee a ke 286
A - Species of the genus Culex occurring in Southeast Asia )
but mot Rnown trom: Thane «3 oe ei ee a eS 286
B - Present status of the Culex fauna of Thailand ............ 289
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. IL.
THE GENUS CULEX IN THAILAND (DIPTERA: i CULICIDAE)!
By
Ralph A. Bram
INTRODUCTION
Background
The culicine mosquito fauna of Thailand has received only scant and
sporadic interest until recent years. Stanton (1920) recorded species oc-
curing commonly in Bangkok; Edwards (1922) included a few general records
from Thailand; Barraud and Christophers (1931) reported on species collected
by J. A. Sinton during a 2 week railway tour of the country; Causey (1937)
summarized information collected during a 4 year stay in Thailand; and
Iyengar (1953) listed 48 species and subspecies from southern areas of Thai-
land.
More recently, Thurman and Thurman (1955) reported on the first
operation of a mosquito light trap in northern Thailand (at Chiang Mai) and
found that 78 percent of species collected were members of the genus Culex.
The most recent faunal list which included species of the genus Culex of Thai-
land was that of E. Thurman, originally presented before the IX Pacific Sci-
ence Congress in 1957 and subsequently published in 1959 and again, with
slight variation, in 1963. This report dealt with species in the northern re-
gion of Thailand (Chiang Mai, Chiang Rai, Lampang, Lamphun, Nan, Phare,
and Tak) but, unfortunately, locality data were not included for records of
species belonging to the genus Culex. The Thurman collection upon which
these reports were based has been deposited in the U. S. National Museum
and forms an important part of the present study.
Extensive mosquito collecting in Thailand by the U. S. Army Medical
Component, South East Asia Treaty Organization from 1961 through 1966 has
ereatly increased our knowledge of the rich fauna of this Southeast Asian
kingdom. Taxonomic Studies based on these collections include those of
Bram (1966), Peyton and Scanlon (1966), Delfinado (1967), Bram and Rattan-
arithikul (1967), Scanlon and Peyton (1967), and Scanlon, Peyton, and Gould
: This work was supported by Research Contract No. DA-49-193-MD-2672
from the U. S. Army Medical Research and Development Command, Office
of the Surgeon General.
2 South East Asia Mosquito Project, Smithsonian Institution, Washington,
D. C. 20560.
2 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
(1967a and 1967b). In the overall study of the genus Culex in Southeast Asia,
Thailand was selected as the starting point because of its central position
within the area under consideration and, most importantly, because the
SEATO collections from Thailand are undoubtedly the most complete and ex-
tensive of any comparable portion of Southeast Asia. The SEATO collections
are now housed in the U. S. National Museum and at this point, at least, have
formed the foundation upon which future studies of the genus Culex will be
based. This paper includes all material received before February 15, 1967.
In this study, 60 species and subspecies belonging to 7 subgenera of
the genus Culex are recognized from Thailand. Of these, 5 species are de-
scribed as new and 19 species are recorded for the first time from Thailand;
7 species previously recorded from Thailand are dismissed as misidentifica-
tions and 6 previous records are regarded as doubtful. In addition, 7 names
are placed in synonymy and 1 subspecies is elevated to species rank.
Format and Treatment
The format utilized for the description of each species requires a
brief explanation. In the synonymy under each species heading, all refer-
ences considered to be pertinent to the understanding of the species are in-
cluded; the mere inclusion of a species name in a checklist or key does not
usually warrant citation, unless there is an historical or taxonomic signifi-
cance. Within the parentheses following each citation, the symbols ¢, ?, L,
and P indicate that the publication deals with at least some part of the male,
female, larva, or pupa respectively; a single asterisk following the symbol
indicates that at least some portion of that stage or sex is illustrated. Ab-
breviations in the synonymy conform to the World List of Scientific Periodi-
cals, 3rd ed., Academic Press, 1952. |
Following the synonymy is a brief diagnosis of the species. A de-
scription of the salient features of the adult female, male, and fourth stage
larva follows the diagnosis. Illustrations of the salient features of each
species are included within the text following the respective species descrip-
tions. The figures have usually been prepared from several specimens and
an attempt has been made to illustrate the most common condition within the
range of variation of each structure. Terminology conforms principally to
that of Belkin (1962); terminology applied to the male antenna and terminalia
of the subgenus Lophoceraomyia follows Colless (1965), except that the divi-
sions of the antennal flagellum are termed flagellomeres and flagellomere I
of this study corresponds to antennal segment II of Colless. Although the
adult females of all but 4 of the species are described, this is the most dif-
ficult stage available for identification. In the subgenera Neoculex, Moch-
thogenes, Lophoceraomyia, and the vishnui subgroup of the sz/zens group in
the subgenus Culex, conclusive determinations cannot usually be based ex-
clusively on the female. Variation within species and the absence of consis-
tent anatomical or colorational features prevent more than subgeneric place-
ment in many instances. Females of the subgenus Lophoceraomyia are par-
ticularly difficult. Some species of the vishnui subgroup are usually recog-
nizable as adult females, but for the most part, conclusive determinations
must depend on associated larval skins. This is indeed unfortunate since the
principle vector of Japanese encephalitis virus in Southeast Asia,
tritaeniorhynchus, belongs to the vishnui subgroup. In the case
of tritaeniorhynchus, the anatomy of the buccopharyngeal armature
Bram: Genus Culex in Thailand 3
has been incorporated as a supplementary character; however, this structure
has not so far been found to be useful in any of the other species of the vishnui
Subgroup. The adult male is described and the terminalia of all but one spe-
cies illustrated. The adult male habitus is usually very similar to that of the
female and only variations from the female are described. In species of the
subgenus Lophoceraomyia, the modified setae on antennal flagellomeres V
through X are described and illustrated. The anatomy of the male terminalia
presents extremely useful features and conclusive determinations can usually
be placed on this structure (or in the case of Lophoceraomyia, in combina-
tion with the antennal flagellomeres). In the subgenera Lutzia and Lopho-
ceraomyia, and the sitiens and bitaeniorhynchus subgroups of Culex, it has
been found desirable to describe and illustrate the lateral aspect of the lateral
plate of the phallosome; in other subgenera, descriptions and illustrations
apply to the dorsal aspect. Species of the vishnuz subgroup exhibit extremely
similar terminalia and determinations based exclusively on the male are not
always reliable. Since the pupal characters have not so far been found taxo-
nomically significant, this stage is not described; however, illustrations of
most pupae of the subgenera Neoculex, Thaiomyia, Culiciomyia, and Culex
have been included. Except for the subgenus Lutzia, the fourth stage larva
presents the most desirable stage for conclusive determinations in the genus
Culex. Whenever possible, an attempt should be made to collect fourth stage
larvae and rear these individually so that skins may be studied concurrently
with the adult males and females. Descriptions and illustrations of the larvae
of all but 3 species of the genus Culex in Thailand have been provided.
Throughout the study, the diagnoses and descriptions of the subgenera and
groups within the subgenera do not necessarily apply to species outside of
Thailand. |
The distribution of each species in Thailand is listed according to
first-order administrative units or changwats, as they are known in Thailand.
Thai locality names conform to the Official Standard Names Gazetteer No. 97
of the U. S. Board of Geographic Names, Washington, D. C., 1966. Names
of changwats are italicized; a double asterisk following a country name indi-
cates a new record for that country. Unless otherwise indicated, distribu-
tion outside of Thailand is in accordance with Stone, Knight, and Starcke
(1959) and Stone (1961 and 1963). Sufficient data are not presently available
to permit critical evaluation of distributional patterns of species within Thai-
land. Figure 1 outlines the 71 changwats of Thailand and indicates the inten-
sity of collection in the various parts of the country. No specimens of the
_ genus Culex have been examined from 22 of the changwats and only 3 or less
species of the genus Culex have been examined from 36 of the changwats.
The area most in need of intensive collecting in respect to the genus Culex is
the Korat Plateau, where only Udon Thani and Nakhon Ratchasima have been
adequately sampled. Valuable distributional data would also be obtained by
more extensive collecting in the Central Valley north of Bangkok.
In the ''Taxonomic Discussion", reasons for proposing nomenclatorial
changes are outlined and discussions of intraspecific variation are presented
where appropriate. Means are also provided for separating a species from
its close relatives. ,
The biologies of the various species of the genus Culex are either un-
known or very poorly understood. With the exception of a few species of the
subgenus Culex and perhaps Lutzia, data are restricted almost exclusively to
4 Contrib. Amer. Ent. Inst., vol..2, no. 1, 1967
a — No Culex studied
SS
ee
\\)
1. Ang Thong
2. Ayutthaya
3. Buriram
4, Chashoengsao
5. Chainat
6. Chaiyaphum
7. Chanthaburi
8. Chiang Mai
9. Chiang Rai
0. Chon Buri
1, Chumphon
Kalasin
Kampong Pate
Kanchanaburi
Khon Kaen
Krabi
Krung Thep (Bangkok)
. Lampang
. Lamphun
Loey
. Lop Buri
Mae Hong Son
. Maha Sarakham
Nakhon Nayok
Nakhon Pathom
. Nakhon Phanom
. Nakhon Ratchasima
. Nakhon Sawan
. Nakhon Si Thammarat
. Nan
. Narathiwat
32. Nong Khai
. Nonthaburi
Pathum Thani
Pattani TL. Yala
ISTHMUS OF
ANDAMAN
SEA
36.
37.
Fig.1
s — 3 or less species of Culex studied
— 4 to 9 species of Culex studied
“41 — 10 or more Species of Culex studied
CAMBODIA
CHANGWATS OF THAILAND
Phetchabun
Phangnga
Phattalung
Phet Buri
Phitsanulok
Phuket
Phichit
. Prachin Buri
Prachuap Khiri Khan
Phare
. Ranong
. Rat Buri
. Rayong
- Roi Et
. Sakon Nakhon
. Samut Prakan
Samut Sakhon
. Samut Songkhram
. Sara Buri
Satun
Sing Buri
. Songkhla
Sisaket
. Sukhothai
Suphan Buri
Surat Thani
Surin
. tak
Thon Buri
<) sePae
Trang
Ubon Ratchathani
Udon Thani
Uthai Thani
Uttar adit
Bram: Genus Culex in Thailand 5
the habitats from which larvae were collected. Surprisingly, our knowledge
of anthropophilic species is almost as incomplete as for the presumed avian
and mammalian feeders. Future studies should certainly concentrate on the
biological aspects of both adult and immature stages.
Zoogeography
Although distributional data for virtually all species of the genus Culex
in Southeast Asia are obviously incomplete and probably in some instances
misleading, a zoogeographical pattern of the mosquito fauna is beginning to
emerge. In the following discussion, the relationships of subgenera in the
Thai fauna with the overall fauna of Southeast Asia will be treated in general
terms.
The 2 species recognized in the subgenus Lutzia from Thailand are
both widely distributed throughout the Oriental region and extend into the
Palearctic region. In the subgenus Neoculex, brevipalpis is widely distributed,
ranging from India and Ceylon eastward and southward through New Guinea
and the Bismarck Archipelago. The other Thai species, tenuipalpis, extends
from the East Himalayas of India into the Shan Highlands of Thailand; Brug's
(1931) record from Java is open to question. The 5 other species of the sub-
genus in Southeast Asia include hayashii, a species with a northern distribu-
tion; sumatranus, recorded from Sumatra and China; simplicicornis, known
only from Borneo; and nematoides and okinawae, both island species.
Of the 3 species of Mochthogenes known from Thailand, malay is
widely distributed, extending from India eastward to Taiwan and southward to
New Guinea; foliatus extends from China southward to the Philippines; and
hinglungensis is known only from Hainan Island and the Shan Highlands of
Thailand. Elsewhere in Southeast Asia, khazani is known from India and
Indochina, and 5 species of the subgenus are endemic to the Philippine Islands.
The single species of the subgenus Acalleomyia is endemic to Malaya, but
future collecting may possibly extend the known distribution at least to the
Isthmus of Kra.
Species of Lophoceraomyia are best considered in respect to the vari-
ous groups and subgroups of the subgenus. The /raudatrix group appears to
be centered in New Guinea but species extend throughout the range of the sub-
genus. Nine species of the group are known from Thailand, and all but 2 spe-
cies, infantulus and rubithoracis, are found in Malaya, Borneo, and/or the
Philippines, but probably do not extend very much north or west of Thailand.
Both infantulus and rubithoracis are widely distributed, extending into India
and China. Species of the brevipalpus subgroup of the mammilifer group were
probably differentiated in the area bordering the South China Sea, and only 2
species, curtipalpis and lucaris, are known to extend into Thailand. Of the
mammilifer subgroup of the mammilifer group, minor, mammilifer, and
bengalensis are widely distributed and range from India to the Philippines or
Malaya; 3 species are known only from Malaya and Thailand; and 6 species
are presently known only from Thailand.
The monotypic subgenus Thaiomyia demonstrates its closest affinity to
Culiciomyia and is presently known only from the Shan Highlands and the Bi-
lank Taung Range in Thailand. Of the 11 species of Culiciomyia known from
Figure 1. A map of Thailand, showing the boundaries of the various
< changwats and general distribution of material studied.
6 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Thailand, bailyz, fragilis, nigropunctatus, pallidothorax, and spathifurca ex-
hibit broad distributions extending, in general, from India through New
Guinea. C. papuensis extends from the Solomon Islands through New Guinea,
Java, the Philippines, and to Thailand, and scanloni ranges from Thailand
through the Philippines and Borneo. Four species, barrinus, spiculothorax,
termi, and thurmanorum, are presently known only from the Shan Highlands
of Thailand. In other areas of Southeast Asia, shebbearei extends from India
to Borneo, but has not been found in Thailand; bahvi is known from Ceylon
with a questionable record from Java; vividiventer is restricted to northern
India, Pakistan, and Nepal; and javanensis and ryukyensis are island species
which have not been reported from mainland Southeast Asia.
In the pipiens group of the subgenus Culex, all 3 species (/uscocephala,
hutchinsoni, and pipiens quinquefasciatus) have very broad distributions which
extend at least from India through Malaya; pipiens quinquefasciatus is, of
course, cosmotropical. The szfzens group of the subgenus Culex is centered
in the Oriental region with some representatives extending into the Australian
and Ethiopian regions. The vishnui subgroup of the sitiens group is composed
of 8 species in Thailand; of these, alienus and perplexus exhibit rather limited
distributions in Malaya and the southern part of the mainland, whereas the
other 6 species are widely distributed and extend from India at least to the
southern part of the mainland (tviiaeniorhynchus being the most widely dis-
tributed). Species of the sitiens subgroup of the sitiens group are restricted
to brackish water habitats of coastal regions. From Thailand veolitoralis
extends through Malaya to New Guinea, and sitiens exhibits an extremely
broad range throughout the Oriental region, Australia, the Pacific islands,
east Africa, and Madagascar. It is surprising that annulirostris, another
member of the sitizens subgroup, has not been found to occur in Thailand; the
2 other species of the subgroup are limited to Pacific islands. Both species
of the gelidus subgroup, gelidus and whitmorei, are widely distributed through-
out the Oriental region and extend even into New Guinea. Three species of
the bitaeniorhynchus subgroup, bitaeniorhynchus, sinensis, and pseudosinensis,
are known from Thailand. The first 2 of these have very broad distributions
throughout the Oriental region and bitaeniorhynchus extends even into Africa
and Madagascar; pseudosinensis is restricted to Malaya, Indochina, and
Thailand. Another species of the bitaeniorhynchus subgroup, geminus, is
known only from Singapore.
In summary, the Culex fauna of Thailand is shared to a large degree
with Malaya and Indonesia, with a small, apparently endemic nucleus in the
subgenera Lophoceraomyia, Thaiomyia, and Culiciomyia, and with a number
of broadly distributed species extending into India. A list of species occur-
ring in Southeast Asia but not known from Thailand may be found as Appendix
A. Appendix B summarizes the status of our knowledge of the Culex fauna of
Thailand.
Bram: Genus Culex in Thailand 7
GENUS CULEX LINNAEUS 1758
Culex Linnaeus 1758, Systema Naturae, 10th ed. 1: 602. Type species: C.
pipiens Linnaeus 1758 (selection of Latreille 1810).
FEMALE. The diagnostic characters are: proboscis not broadened
distally and not sharply recurved, although the apex may be laterally com-
pressed and slightly downcurved; antennal flagellomeres neither short nor
thick; anterior pronotal lobes of normal size and widely separated; posterior
margin of scutellum trilobed, the marginal setae in three groups; postnotum
without scales or setae; spiracular and postspiracular bristles absent; wing
membrane with distinct microtrichia and vein 1A long, reaching wing margin
beyond level of junction of Cul and Cu2; dorsal plume scales not forked apical-
ly; mid and hind femora without distinct tufts of scales at apices; tarsomere I
of fore leg longer than I-V combined, IV longer than wide; pulvilli present on
all legs; claws of hind legs very small and inconspicuous. Additional charac-
ters are as follows. Head. Eyes contiguous, very narrowly separated above
the antennae, with the orbital bristles well developed; length of palpus ap-
proximately 1/4 or less than that of the proboscis; length of antenna equal to
or greater than that of proboscis; decumbent scales of the vertex narrow at
the occiput, becoming somewhat broader at the orbital line, particularly the
posterolateral area; erect scales always forked, usually numerous. Thorax.
Scutum sparsely to heavily clothed with narrow scales, acrostichal bristles
present or absent, dorsocentral bristles well developed; integument of the
pleuron with or without distinctive dark patterns, and with or without scat-
tered patches of dull, pale scales (never densely covered with broad, distinc-
tively ornamented scales); pleural setation varied, present on the anterior
pronotal lobe, posterior pronotal lobe, propleuron, sternopleuron, prealar
area, and upper mesepimeron, those of the lower mesepimeron either present
or absent. Wings. Vein scales variable, but dorsal plume scales never
forked; cell Rg always longer than vein Rg,3. Abdomen. Tergum I with only
a small scale patch in the middle; terga and sterna II-VI uniformly scaled;
apex of abdomen truncate.
MALE. The diagnostic characters are the same as those of the fe-
male in addition to the following features of the terminalia: proctiger strongly
developed and with a transverse crown of numerous apical spines; basimere
with a subapical lobe which normally exhibits strongly modified or specialized
setae; phallosome usually complex. Additional characters are as follows.
Head, Antenna plumose, occasionally equal to or slightly less than the length
of the proboscis; length of the palpus variable (when longer than proboscis, the
last 2 segments usually slender and upturned). Legs. Tarsal claws of the
fore and mid legs enlarged, those of the hind legs as small as in the female.
Terminalia, Basimere well developed, without scales; a well developed sub-
apical lobe always present, frequently separated into a proximal and distal
division; the proximal division typically exhibits 3 strong, subequal rods, and
the distal division generally exhibits several variable setae, one of which is
frequently leaf-shaped; distimere always well developed, although somewhat
variable in shape; phallosome complex, usually consisting of a pair of vari-
able lateral plates.
LARVA. The diagnostic characters are: siphon with a short, movable
dorsal valve present; accessory ventral, ventrolateral, or dorsolateral hairs
8 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
present on the siphon, but without a differentiated pair of subventral hairs at
the extreme base (if the accessory hairs are short and indistinct, then the
siphon is very long); a distinct basolateral acus always present on the siphon;
thoracic hair 13-P always absent. Additional characters are as follows.
Head, Mouthparts usually not modified (except in the predaceous subgenus
Lutzia and the brevipalpus subgroup of the subgenus Lophoceraomyia); hairs
1,4,5,6,7-C well developed; 16,17-C present or absent, when present repre-
sented by small spicules; antenna long and constricted beyond insertion of
hair 1-A (except in Lutzia and the brevipalpus subgroup of Lophoceraomyia);
hairs 2,3-A usually subapical (except in the groups noted above). Thorax.
Integument spiculose or glabrous; hairs 1,2-P always well developed, usually
single; 3-P variable; 4,5, 6,7-P well developed; 8-P variable; 9,10,11,12-P
always inserted on the same tubercle, 12-P the strongest of the group; 14-P
simple, single or bifid. Abdomen. Integument glabrous or spiculose, with-
out sclerotized plates except for the saddle on segment X; hair 1-I-VIII never
palmate; comb always present on segment VIII but variable, never with a
common sclerotized plate at the base of the scales; pecten variable, absent
only in C. (Thaiomyia) dispectus; 3 or more pairs of subventral tufts usually
inserted beyond the pecten; segment X with the saddle fomplete except in C,
(Culex) neolitoralis; ventral brush with at least 8 tufts’, most of which are
inserted on the grid; 2 pairs of anal gills variously developed.
PUPA. So far, features have not been found which permit a definitive
characterization of the genus. Some of the salient features are as follows.
Cephalothorax. All hairs present, with variable development; 6-C weak,
Single or branched, usually distinctly weaker and shorter than 7-C; trumpet
of variable length and form, but always with a pinna and never placed ona
distinct tubercle. Abdomen. All normal hairs usually present; 9-IV-VI
varied in position but always removed from caudolateral angle of tergite,
sometimes spiniform but never on all segments; 1-IX usually present, always
a Simple bristle; 1-X absent; paddle variously developed but never narrow;
midrib usually strongly developed and dividing paddle into subequal parts;
1,2-P usually both present, sometimes one or both hairs absent.
i DISTRIBUTION. The genus Culex is worldwide in distribution, with
marked concentration in the tropical regions. Representatives of the genus
are found throughout the Oriental region and have been collected in virtually
every changwat of Thailand. Of the 17 subgenera recognized for the genus,
only 8 have been recorded from the Oriental region and of these, 7 are known
from Thailand (Acalleomyia has been recorded only from Malaya, but may be
present on the Isthmus of Kra in Thailand).
TAXONOMIC DISCUSSION. Within the genus Culex, all subgenera
cannot be separated in every instance exclusively on the basis of the adult
female (particularly when dealing with individuals of the subgenera Lopho-
ceraomyia, Culiciomyia, Neoculex, and Thaiomyia) and the pupae have not
been found to exhibit consistent subgeneric features. In these instances it is
necessary to have not only the adult males and females, but their associated
: These tufts are often referred to as occurring in pairs whereas in actual
fact they are staggered in zigzag fashion down the grid, each bar of the grid
having 1 tuft only. Here, as elsewhere, the figure refers to the total number
of tufts and not to pairs.
Bram: Genus Culex in Thailand 8)
larval skins as well.
: BIOLOGY. Eggs of Culex are typically laid in rafts on various water
surfaces and there is no evidence that eggs of any species are resistant to
desiccation. Larval habitats include a wide variety of ground water accumu-
lations (primarily fresh, but also brackish) as well as tree holes, bamboo
internodes and stumps, pitcher plants, and containers. Members of the sub-
genus Lutzia are predaceous on other mosquito larvae and members of the
brevipalpus subgroup of the subgenus Lophoceraomyia are apparently preda-
ceous on other small arthropods which accumulate in their pitcher plant envi-
ronments. Adults of the subgenus Culex probably feed to a large degree on
mammals and birds, while those of Lophoceraomyia and Lutzia are probably
primarily avian feeders, and those of Neoculex are possibly reptilian and am-
phibian feeders. Feeding behavior of the other subgenera are even less well
known.
Members of the genus Culex have been incriminated in the transmission
of a number of pathogenic organisms in Southeast Asia, but most of these are
probably incidental infections or isolations. Three species, all members of
the subgenus Culex, are currently recognized as actual or potential disease
vectors of considerable importance in Thailand. Two species, gelidus and
tritaeniorhynchus, are important vectors of Japanese encephalitis in South-
east Asia, and pipiens quinquefasciatus is the most important vector of ur-
ban filariasis caused by periodic Wuchereria bancrofti (a rare disease in
Thailand, but very common in neighboring countries, such as Burma).
Species of the genus Culex are susceptible to a wide variety of patho-
gens and parasites and are prey for a great number of predators. Jenkins
(1964) reviewed the organisms attacking medically important arthropods and
compiled an impressive list of pathogens and parasites of Culex. Included in
this listing were viruses (2 species), rickettsiae (3 species), bacteria (5
species), spirochaetes (2 species), fungi (20 species), protozoa (41 species),
rotatoria (2 species), nematodes (8 species), and trematodes (3 species).
Not all species listed cause mortality and those that do are generally restricted
to the polyhedrosis viruses, a few species of bacteria, several of the fungi
and fungi imperfecti (particularly the Coelomomyces), some protozoa, and
mermithid nematodes. Most isolations have been made from the mosquito
larva, but some have been made from adults.
KEY TO THE SUBGENERA OF CULEX IN THAILAND - ADULTS
ae Four or more strong, lower mesepimeral bristles present; relatively
large species (wing length usually ranging from 4to5 mm)......
Lutzia (p. 11)
Mesepimeral bristles absent, or if present, represented by 1 or 2
rather weak ones; in general moderately sized to small species
(wing. length uswally lese tian 4mm) 5006 ee a ee: 2
2(1). Pleuron with distinct scale patches at least on the upper and lower
sternopleuron and the anterior meSepimeron...... Culex (p. 163)
Pleuron without distinet scale patches... eee ee 3
3(2). Acrostichal bristles well developed on the scutum; palpus of the male
10
4(3).
5(4).
6(5).
2(1).
3(2).
4(3).
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
less than 1/4 the length of the proboscis. . . .Mochthogenes (p. 33)
Acrostichal bristles not developed except at extreme anterior end and
rarely near the prescutellar space; palpus of the male greater
than’ 1/4 the Jength of the proboscis! 6°) cee aes owes 4
Scaling of the scutum sparse, rough in appearance; male antenna with
specialized scales and setae usually present on flagellomeres V-_.
IX, always present on Vil and VII...... Lophoceraomyia (p. 42)
Scaling of the scutum very dense, Smooth in appearance; male
antenna without specialized scales or setae .............0-. 5)
Palpus of the male approximately 1/2 to 3/4 the length of the
Sronoecin ec 6c ol ee OE Os es a es Neoculex (p. 23)
Palpus of the male greater than or approximately equal to the length
OD Tie OP OG Cte i os I Se OR OC ie ORO ws 6
Palpal segment III of the male with a characteristic ventrolateral,
linear series of lanceolate scales......... Culiciomyia (p. 117)
Palpal segment III of the male without lanceolate scales..........
Thaiomyia (1 species, dispectus, p. 113)
KEY TO THE SUBGENERA OF CULEX IN THAILAND -
FOURTH STAGE LARVAE
Pecten extending nearly to the apex of the siphon; mouthparts
adapted for predation; antennal hair 1-A represented by a short,
single, simple seta inserted on the proximal half of the shaft;
wery Inree Gpeciog o.oo US ie eens a Lutzia (p. 11)
Pecten restricted to the basal half of the siphon or absent; mouth-
parts not adapted for predation (or if moderately so, other
characters not as above); antennal hair 1-A generally multiple,
always inserted on the distal half of the shaft; small to moderately
POG BOCs i Cn A, as ee a, a Pe eS 2
Pecten absent.......... Thaiomyia (1 species, dispectus, p. 113)
Pecten represented by 2 or more distinct teeth............... 3
Ventral brush consisting of 8 hair tufts inserted on the grid; head
hair 1-C fine and filamentous and thoracic hair 3-P much
slenderer than 1-P and usually about half its length, ov if head
hair 1-C stout, 1-P bifid or trifid and 3-P with 2 or more
Gratiches cece ip ea GL PS PUNO WOES Culiciomyia (p. 117)
Ventral brush consisting of 10 or more hair tufts; if head hair 1-C
is filamentous, then thoracic hairs 1,3-P single and of the same
order of length and thickness, ov 1-C robust and 1-P un-
POPU ee ee Pe ES ae a a
Thoracic hair 3-P always single, of the same order of length and
Phichkness ae t+ Be Ge eae ea Bie Mee Culex (p. 163)
Thoracic hair 3-P single or branched, slenderer than 1-P and usually
Bram: Genus Culex in Thailand 11
less than half lis lengths a Be Ee 3)
5(4). Siphon shorter than the anal segment; antennal hair 1-A very weak,
simple; mouthparts adapted for predation................+..
Lophoceraomyia (brevipalpus subgroup, p. 75)
Siphon longer than the anal segment; antennal hair 1-A strong,
multiple, pectinate; mouthparts not adapted for predation...... 6
6(5). Head hair 14-C single, often dendritic beyond the basal half.......
Neoculex (p. 23)
Head hair 14-C bifid, with the branches strongly divergent from the
7(6). Individual pecten tooth fringed with many fine, parallel, closely placed
denticles distally and a few coarse, widely placed denticles prox-
imally; head hairs 5,6-C short, weak, 5-C about 1/2 the length
GE BES OPEL Re POG a es Mochthogenes (p. 33)
Individual pecten tooth without 2 distinct types of denticles as in-
dicated above (although the proximal 2 or 3 denticles may be en-
larged); head hairs 5, 6-C strong, long, subequal in length......
Lophoceraomyia (in part, p. 42)
SUBGENUS LUTZIA THEOBALD 1903
Lutzia Theobald 1903, Mon. Cul. 3: 155. Type species: Culex bigoti
Bellardi 1862.
Jamesia Christophers 1906, Sci. Mem. med. sanit. Dep. India 25: 12 (pre-
occupied by Jeckel 1861). Type species: Culex concolor Robineau-
Desvoidy 1829 (selection of Edwards 1932, in Wytsman, Genera
Insect., fasc. 194: 190, as Culex fuscanus Wiedemann).
The adult female may be recognized by its relatively large size (wing
length approximately 4 to 5 mm), the ventral banding of the proboscis, and
the numerous, strong lower mesepimeral bristles. The male may be recog-
nized by the above mentioned characters as well as the simple phallosome of
the terminalia and the strong but reduced number of setae on the subapical
lobe of the basimere. The fourth stage larva may be identified by its large
size, the predaceous mouthparts, and the pecten which extends nearly to the
apex of the siphon.
FEMALE. Very large species for the genus with conspicuous orna-
mentation. Head. Proboscis with a pale median band of scales on the ventral
surface and frequently encompassing the proboscis; antenna longer than the
proboscis; torus with a number of broad scales on the inner margin. Thorax.
Scutum with very dense scaling, usually with distinct but variable patches of
light scales; acrostichal bristles well developed, but rather short; distinct
pleural scale patches present; 4 or more strong, lower mesepimeral bristles
present.
MALE. Head, Palpus and antenna longer than the proboscis; palpal
segments IV and V upturned. Terminalia. Subapical lobe of the basimere
with strongly developed setae, but without a leaflike seta; distimere normal
12 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
for the genus; phallosome simple; proctiger crowned with a dense tuft of
rather slender, short spicules; usually 5 or more short cercal setae present.
LARVA. Head, Head capsule elongate; antenna glabrous, gradually
tapering to a truncate apex, and not constricted beyond the insertion of hair
1-A which is short, single, simple and inserted on the proximal half of the
shaft; head hair 1-C single, simple; 4-C dendritic with about 4 or 5 branches;
5,6-C single, simple, very strong and long; 16,17-C absent; mentum with
very strong teeth; mouthparts adapted for predation and mouth brushes very
broad and lamellate; integument of the head covered with a conspicuous pat-
tern of minute spicules. Thorax. Integument covered with a conspicuous
pattern of very dense, small, sharply pointed spicules; prothoracic hairs |
relatively short. Abdomen. Integument covered with spicules which are not
nearly as apparent as those of the thorax; comb consisting of a broadly tri-
angular patch of fan-shaped scales; siphon variable, index approaching 1:1,
covered with a dense conspicuous pattern of sharply pointed spicules which
seem to be modified in the form of reticulose platelets; approximately 10 sub-
ventral tufts inserted on the siphon; pecten extending almost to the apex; sad-
dle completely ringing the anal segment; ventral brush consisting of approxi-
mately 15 individual tufts inserted within the grid.
PUPA. Very large, but similar in chaetotaxy to those of the subgenus
Culex. Paddle broad.
DISTRIBUTION. Members of the subgenus Lutzia are found in tropical
and semitropical areas of both the Old and New World. Both species in Thai-
land extend throughout the Oriental region. C. halifaxii reaches through the
Australian region to the Solomon Islands and northward to Japan, China, Korea,
the U.S.S.R. and India, whereas fuscanus extends through Taiwan and Japan
to Korea, parts of China, the Maritime Province of U.S.S.R. and India.
TAXONOMIC DISCUSSION. Belkin (1962) suggested that Lutzia is an
ancient derivative of the genus Culex with strongest affinities to the subgenera
Culex and Culiciomyia. Edwards (1932) recognized two groups within the
subgenus Lutzia, restricted respectively to the Old and New Worlds. It now
appears that the Old World group (Jamesia) is represented by 3 species:
tigripes Grandpre and Charmoy in the Ethiopian region, and fuscanus and
halifaxii in other Old World areas. Larvae of the species found in Thailand
are indistinguishable and adults are extremely variable. The male termi-
nalia, however, exhibit consistent differences in the phallosome which are
considered to be diagnostic.
BIOLOGY. Larvae of the members of the subgenus Lutzia are preda-
ceous, principally on mosquito larvae of other species. They are found most
often in ground water habitats, frequently where there is a high organic con-
tent, in association withC, pipiens quinquefasciatus. Adult females probably
feed, to a great extent, on avian hosts and seldom attack man.
KEY TO SPECIES OF THE SUBGENUS LUTZIA IN THAILAND -
ADULT FEMALES
Abdominal terga V-VII entirely pale scaled, or with very broad
apical pale bands; terga I-IV entirely dark, or with very
narrow pale apical bands; median pale band of the proboscis
broad, clearly encircling if 2645.6... o6'ae bea fuscanus (p. 13)
Abdominal terga entirely dark, or all with pale apical bands of ap-
Bram: Genus Culex in Thailand 13
proximately the same width; median pale band of the proboscis
narrow, most prominent on the ventral surface. . . halifaxii (p. 17)
KEY TO SPECIES OF THE SUBGENUS LUTZIA IN THAILAND -
MALE TERMINALIA
Phallosome, when observed in lateral aspect, without prominent
denticles extending from the lateral plate; however, faint stria-
tions or suggestions of teeth may be observed. . . . fuscanus (p. 13)
Phallosome, when observed in lateral aspect, with a series of up to
10 prominent denticles extending from the internal margin of the
1eier al VIALE Ci as eal ’s 2 PA a ee a halifaxii (p. 17)
(The immature stages of the subgenus Lutzia cannot be distinguished at the
present time)
CULEX (LUTZIA) FUSCANUS WIEDEMANN 1820
(Figures 2 and 4A)
Culex fuscanus Wiedemann 1820, Dipt. Exot. 1: 9 (<, 9).
Culex concolor Robineau-Desvoidy 1827, Mem. Soc. Hist. nat. Paris 3: 405
(*); Edwards 1922, Indian J. med. Res. 10: 275 (synonymy).
Culex setulosus Doleschall 1875, Natuurk. Tijdschr. Ned. -Ind. 14: 384 (¢,
2*); Edwards 1922, Indian J. med. Res. 10: 275 (synonymy). ,
Culex luridus Doleschall 1875, Natuurk. Tijdschr. Ned. -Ind. 14: 384 (o*, ?*);
Barraud 1934, Fauna Brit. India, Diptera 5: 341 (synonymy).
Lutzia fuscanus (Wiedemann): Edwards 1922, Indian J. med. Res. 10: 275
(**); Barraud 1924, Indian J. med. Res. 11: 973 (o*, 9).
Culex (Lutzia) fuscanus Wiedemann: Edwards 1932, in Wytsman, Genera
Insect. , fasc. 194: 191 (taxonomy); Barraud 1934, Fauna Brit. India,
Diptera 5: 341 (o*, ?, L); Li and Wu 1935, Yearb. Bur. Ent. Chekiang
4: 98 (L*); Bonne-Wepster and Brug 1939, Geneesk. Tijdschr. Ned. -
Ind. 79: 1261 (L*); Bohart 1945, Navmed 580: 72 (o*); T'an 1945,
Sinensia 16: 46 (L*); Bonne-Wepster 1954, Roy. trop. Inst. Amst.
Spec. Pub. 111: 107 (oc, 9*, L*); Bohart 1956, Ins. Micronesia 12: 75
(o*, 2); Ma and Feng 1956, Acta ent. Sinica 6: 172 (L*); Ikeshoji 1966,
Jap. J. exp. Med. 36: 321-334 (biology); Delfinado 1966, Mem. Amer.
ent. Inst. 7: 97 (o'*, 9, L*, P*).
Color patterns of the adult female are extremely variable, particularly
on the scutum, the legs, and the abdomen; however, abdominal terga I-IV usually
exhibit very narrow apical pale bands and terga V-VIII are totally pale. In the
male the phallosome is distinctive, being robust, but very simple and lacking
prominent denticles. The larva cannot be distinguished from that of halifaxii.
FEMALE. (Figure 2). In general a large species for the genus Culex
with outstanding color patterns particularly on the abdomen and scutum. Head.
(Figure 2B). Proboscis dark brown, usually with an extremely broad median
band of light scales clearly encircling the proboscis and with scattered pale
scales proximal to the band; palpus similar in color to the proboscis, with white
14
Contrib. Amer. Ent. Inst., vol. 2,
C. (Lutzia) fuscanus
no. 1, 1967
Bram: Genus Culex in Thailand 15
scales scattered among the dark particularly on the apex and on the internal
margin; decumbent scales of the vertex uniformly pale; erect scales numerous,
uniformly dark brown. Thorax. (Figure 2B, E). Scutum with scales forming
distinct, but variable dark and light patterns; the most prominent light scale
patches present on the anterior acrostichal area, the humeral area, the pos-
terior fossal area, and around the prescutellar space; integument of the pleuron
light brown; distinct scale patches present on the prealar area, the upper
sternopleuron, the posterior sternopleuron, and the upper mesepimeron;
bristles also present in these areas in addition to 3 to 7 strong lower mesep-
imeral bristles. Wing. (Figure 2C). Legs. (Figure 2D). Anterior surface
of the hind femur with a broad median stripe of pale scales and with a distinct
knee spot at the apex; hind tibia with numerous white scales which also form
somewhat of a median stripe; hind tarsomere I mostly pale, tarsomeres II-V
dark but with a variable amount of pale scaling; markings of the fore and mid
legs similar to the hind legs. Abdomen. (Figure 2F). Terga I-IV dark,
frequently with narrow pale apical bands or with scattered pale scales, terga
V-VIII completely pale or with broad, pale apical bands; sterna uniformly
pale.
MALE. Similar in general appearance to the female, but frequently
with the pale areas less prominent or slightly modified. Head, (Figure 2A).
Median pale band of the proboscis usually modified to be more distal and less
distinct than in the female; palpus dark, with variable pale scaling on seg-
ments I-V. Terminalia. (Figure 4A). Basimere conical, without scales;
subapical lobe of the basimere with the basal rods strong, hooked distally;
beyond the rods are approximately 5 short, fine setae, none of which are leaf-
shaped; distimere robust, with up to 7 fine, scattered setae; phallosome sim-
ple, consisting of a gently curved lateral plate, the internal surface exhibiting
- a series of up to 7 fine denticles or striations which do not protrude when
observed in lateral aspect; proctiger well developed, crowned with a dense
tuft of strong spines; cercal setae variable, approximately 8 in number; basal
sternal process well developed, straight.
LARVA. The larva of fuscanus cannot be distinguished from that of
halifaxii; for a discussion of structure and chaetotaxy, see under the latter
Species.
TYPE DATA. Syntypes of fuscanus from India are in the Naturhist-
orisches Museum, Vienna, Austria. No type locality was designated for
concolor and the type specimen is non-existent. Syntypes of setulosus from
Gombong, Java are in the Naturhistorisches Museum, Vienna, Austria. The
type locality of /uridus was designated as Gombong, Java but the location of
the type specimen is unknown.
DISTRIBUTION. This species is probably distributed throughout THAI-
LAND. During this study specimens have been examined from: Chiang Mai,
Chon Buri, Krung Thep, Nakhon Ratchasima, Phet Buri, Prachuap Khiri
Khan, Rayong, Surat Thani, Thon Buri, and Udon Thani. It has also been
Figure 2. C. (Lutzia) fuscanus. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax; C,
<—_- dorsal aspect of the female wing; D, anterior surface of the fe-
male legs; E, dorsal aspect of the female scutum and scutellum;
F, dorsal aspect of the female abdomen.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
16
C. (Lutzia) haiifaxti
Bram: Genus Culex in Thailand 17
recorded from the following additional countries: INDIA, CEYLON, ANDA-
MAN ISLANDS, BURMA, MALAYA, SUMATRA, JAVA, BORNEO, PHILIP-
PINES, INDOCHINA, CHINA, TAIWAN, U.S.S.R. (Maritime Province),
KOREA, JAPAN, CAROLINE ISLANDS, and PALAU ISLANDS.
The following specimens have been examined during this study: 46
males (10 with associated larval and pupal skins) and 68 females (16 with as-
sociated larval and pupal skins) from Thailand as well as 184 males and 295
females from other areas within its range. Numerous whole larvae were
examined, but distinction could not be made between species of the subgenus.
BIOLOGY. The bionomics of this species has recently been studied in
Rangoon, Burma by Ikeshoji (1966a). It was found that oviposition preferences
of the females correspond very closely to those of pipiens quinquefasciatus,
although the latter species is capable of tolerating higher levels of pollution.
Upon hatching, the first instar larva immediately attacks another larva of
Similar size and over the whole period of development the individual will con-
sume an average of 363.8 larvae, most of which are in the early instars. Al-
though the association of fuscanus larvae with those of pipiens quinquefasciatus
was very striking, the population density of fuscanus remained relatively low
throughout the year, averaging 21 larvae and pupae of fuscanus per 10,000 to
00,000 pipiens quinquefasciatus larvae even when both populations were at
their peaks during the January-February period. Adult females apparently
feed primarily on avian hosts and could not be induced to feed on man. Ikeshoji
concluded that although fuscanus larvae are potent destroyers of pipiens
quinquefasciatus larvae under field conditions, the predator persists at such
a low level that no significant controlling effect is registered on the pipiens
quinquefasciatus population .
CULEX (LUTZIA) HALIFAXII THEOBALD 1903
(Figures 3, 4B, and 5)
Culex halifaxii Theobald 1903, Mon. Cul. 3: 231 (¢); Edwards 1913, Bull. ent.
Res. 4: 234 (taxonomy).
Culex multimaculosus Leicester 1908, Cul. Malaya: 155 (, 9); Edwards 1913,
Bull. ent. Res. 4: 234 (synonymy).
Culex aureopunctis Ludlow 1910, Canad. Ent. 42: 195 (?); Edwards 1922,
Indian J. med. Res. 10: 470 (synonymy).
Lutzia vorax Edwards 1921, Bull. ent. Res. 12: 327 (o*); Barraud 1924,
Indian J. med. Res. 11: 975 (o*,?). NEW SYNONYMY .
Lutzia halifaxi (Theobald): Edwards 1921, Bull. ent. Res. 12: 327 (co);
Edwards 1924, Bull. ent. Res. 14: 391 (distribution); Barraud 1924,
Indian J. med. Res. 11: 975 (o'*, 9); Paine and Edwards 1929, Bull.
ent. Res. 20: 307 (distribution).
Lutzia raptor Edwards 1922, Indian J. med. Res. 10: 275 (o*); Barraud 1924,
Indian J. med. Res. 11: 974 (*o*). NEW SYNONYMY.
Culex (Lutzia) halifaxi(i) Theobald: Edwards 1932, in Wytsman, Genera
Figure 3. C. (Lutzia) halifaxi. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax;
a C, dorsal aspect of the female scutum and scutellum; D,
anterior surface of the female legs; E, dorsal aspect of the fe-
male wing; F,G,H, andI, dorsal aspect of variations in the
female abdomen. ,
18 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Insect. fasc. 194: 191 (taxonomy); Barraud 1934, Fauna Brit. India,
Diptera 5: 344 (o'*, 9, L*); Bonne-Wepster and Brug 1937, Geneesk.
Tijdschr. Ned. -Ind. 77: 62 (o,?); Bonne-Wepster and Brug 1939,
Geneesk. Tijdschr. Ned. -Ind. 79: 1263 (L); Knight, Bohart and
Bohart 1944, Nat. Res. Counc. Div. med. Sci. 45 (key); Bohart 1945
Navmed 580: 72 (c'); Laird 1947, Trans. and Proc. Roy. Soc. N. Z.
76: 459 (L*); Knight and Chamberlain 1948, Proc. helm. Soc. Wash.
15: 18 (P*); Perry 1949, Amer. J. trop. Med. 29: 754 (L*); Penn
1949, Pacif. Sci. 3: 68 (P*); Bick 1951, Pacif. Sci. 5: 415 (ecology);
Bonne-Wepster 1954, Roy. trop. Inst. Amst. Spec. Pub. 111: 108
(o', 2, L); Belkin 1962, Mosq. S. Pacif. 1: 220 (o'*, 9, L*, P*); Assem
and Bonne-Wepster 1964, Zool. Bijdr. (Leiden) 6: 109 (o, 9);
Delfinado 1966, Mem. Amer. ent. Inst. 7: 99 (c*, 9).
Culex (Lutzia) raptor (Edwards): Edwards 1932, in Wytsman, Genera Insect.
fasc. 194: 191 (taxonomy); Barraud 1934, Fauna Brit. India, Diptera
5: 343 (o*, 2, L).
Culex (Lutzia) vovax (Edwards): Edwards 1932, in Wytsman, Genera Insect.
fasc. 194: 191 (taxonomy); Barraud 1934, Fauna Brit. India, Diptera
5: 344 (o*, 9, L); Robertson and Hu, 1934, China J. 20: 356 (distribu-
tion); Hsiao and Bohart 1946, Navmed 1095: 24 (distribution); Bohart
and Ingram 1946, Navmed 1055: 70 (o*, 9, L*); LaCasse and Yamaguti
1950, Mosq. Fauna Japan and Korea: 251 (o*, 9*, P*, L*); Monchadskii
1951, Moscow Zool. Akad. Nauk SSSR 37: 242 (L*); Asanuma and
Nakagawa 1953, Misc. Rep. Res. Inst. nat. Resour. 31: 91 (P*);
Bohart 1956, Ins. Micronesia 12: 73 (o*,?, L*); Ma and Feng 1956,
Acta ent. Sinica 6: 172 (L*); Hara 1957, Jap. J. exp. Med. 27: 52
(2*); Lien 1962, Pacif. Ins. 4: 630 (distribution).
Color patterns of the adult female are even more variable than in
fuscanus; however, the abdominal banding patterns and the incomplete pale
band of the proboscis are diagnostic. The phallosome of the male is charac-
terized by having 3 to 10 strong denticles which project prominently from the
inner margin of the lateral plate. The larva cannot be distinguished from that
of fuscanus.
FEMALE. A large species for the genus Culex with prominent colora-
tional features on the legs, scutum, and abdomen. Head. (Figure 3B).
Proboscis dark brown with a very narrow, but variable ventral median band
of pale scales which occasionally encircle the proboscis; palpus similar in
color to the proboscis, with or without white scales scattered among the dark;
decumbent scales of the vertex uniformly creamy white; erect scales usually
lighter than those on the rest of the vertex. Thorax. (Figure 3B,C). Scutum
basically covered with bronze-brown scales, but with a variable pattern of
silver or white scales on the anterior promontory, the humeral area, and
scattered in and around the prescutellar space; acrostichal bristles numerous
Figure 4. Dorsal view of the male terminalia with inserts of the lateral
aspect of the phallosome: A, C. (Lutzia) fuscanus; B, C. (Lutzia)
halifaxit.
—_>
Bram: Genus Culex in Thailand
rH UP iN a
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radars “Nt
“epee | FT Mth NPP ratty,
ie
ca
Vyt yy
C.(Lutzia) halifaxii
ht pes
19
20 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
anteriad of the scutal angle; integument of the pleuron uniformly light brown;
distinct white scale patches on the prealar area, the upper sternopleuron,
the posterior sternopleuron, the upper mesepimeron, and the anterior
mesepimeron; bristles also present in these areas in addition to several strong
lower mesepimeral bristles. Wing. (Figure 3E). Legs. (Figure 3D). The
color patterns of all legs similar; femora basically dark brown with flecks of
white scales scattered to a greater or lesser degree and with a narrow, pale ©
apical band; each tibia also basically dark brown, but with the light scales
forming somewhat of a longitudinal stripe and with a narrow apical band; tarsi
light brown with very few randomly placed light scales; considerable variation
exists in the pattern of light scales on the femora and tibiae. Abdomen. Varia-
tion in tergal banding patterns illustrated in figure 3F-1; in the typical halifaxii
form (31), all of the terga are dark scaled without light basal or apical bands,
however, tergum VII has a basal triangular spot of light scales laterally and
VI has a narrow white basal band; another condition exists in which all terga
exhibit moderately broad apical white bands (3F) and conditions intermediate
to the 2 extremes are common (3G, H); sterna with variable patterns of mixed
dark and light scales.
MALE. Similar in general appearance to the female. Head. (Figure
3A). Proboscis with the median pale band always present on both the dorsal
and ventral surfaces; palpus usually with scattered pale scales on segments
II-V. Terminalia. (Figure 4B). Very similar to fuscanus; the phallosome,
when viewed in lateral aspect, with a series of up to 10 prominent denticles
protruding from the inner margin of the lateral plate.
LARVA. (Figure 5). Head. Integument rather darkly pigmented,
particularly around the insertions of hairs 5,6-C and covered with a uniform,
dense pattern of small, tooth-like spicules; mouthparts adapted for predation
and mouth brushes very broad and lamellate; antenna gradually tapering to a
truncate apex, not sharply constricted beyond insertion of hair 1-A which is
represented by a single, simple seta inserted on the proximal half of the
shaft; hairs 2,3-A single, simple, subequal in length, inserted somewhat
below the apex of the shaft; head hair 1-C single, simple, arising from a
distinct, small protruberance; 4-C dendritic, with 4 or 5 branches; 5, 6-C
single, simple, very strong and long; 16,17-C absent. Thorax. Integument
covered with a conspicuous pattern of very dense, small, sharply pointed
spicules; thoracic hairs 1-7-P all single, simple, rather short; 8-P branched,
pectinate; 14-P single, simple. Abdomen. Integument minutely spiculose,
the spicules not nearly as apparent as those of the thorax; comb consisting of
approximately 35 fan-shaped scales arranged in a broad, triangular patch;
siphon index ranging from 1:1 to 1:1.5; the siphon covered with a dense,
conspicuous pattern of sharply pointed spicules which are modified in the form
of reticulose platelets; approximately 12 to 20 tufts of pectinate setae inserted
in a straight, single midventral line and a pair of small, bifid, simple tufts
inserted dorsad of the pecten; pecten consisting of from 9 to 12 teeth located
throughout the length of the siphon except at the extreme base and apex;
individual pecten tooth consisting of a very strong apical spine and 1 or 2
Figure 5. C. (Lutzia) halifaxii. Fourth stage larva: Dorsoventral view of
the head and prothorax, and lateral aspect of the terminal ab-
dominal segments.
—>>-
21
Genus Culex in Thailand
e
e
Bram
E
E
°
I L. HWANG
C. (Lutzia) halifaxii
22 Contrib. Amer. Ent. Inst., vol. 2. no. 1, 1967
(usually 1) strong basal barbs; ventral brush consisting of from 14 to 16 in-
dividual tufts inserted within the grid; anal gills very short; anal saddle
complete, spiculose.
TYPE DATA. The holotype female of halifaxii from Dindings, Malaya
is in the British Museum. Lectotype male of multimaculosus is hereby
designated: Cotype male (terminalia mounted and attached to pin), ''Kuala
Lumpur, Fed. Malay States, Dr. G. F. Leicester, 1912-350", in the British .
Museum. The holotype female of aureopunctis from Cotabato, Mindanao,
Philippines is in the U. S. National Museum. The holotype male (terminalia
mounted and attached to pin) of raptor from Amritsar, India is in the British
Museum. The holotype male (terminalia mounted and attached to pin) of
vorax from Tokyo, Japan is in the British Museum.
DISTRIBUTION. This species is undoubtedly distributed throughout
THAILAND and has been examined from: Chanthaburi, Chiang Mai, Chon
Buri, Kanchanaburi, Krung Thep, Lampang, Nakhon Nayok, Nakhon Ratchasima,
Narathiwat, Nonthaburi, Phattalung, Phet Buri, Prachuap Khiri Khan, Ranong,
Songkhla, Thon Buri, Trang, and Udon Thani. C. halifaxii, as presently
recognized to include vaptor and vovax, has been reported from:MALAYA,
INDIA, CEYLON, CHINA, INDONESIA, PHILIPPINES, NEW GUINEA,
BISMARCK ARCHIPELAGO, SOLOMON ISLANDS, AUSTRALIA, BURMA,
INDOCHINA, JAPAN, U.S.S.R. (Maritime Province), KOREA, OGASAWARA
GUNTO, RYUKYU-RETTO, TAIWAN, MARIANA ISLANDS, and NEPAL.
During this study 196 adult females and 114 adult males (33 of which
had associated larval and pupal skins) were examined from Thailand, as well
as 138 specimens from other countries.
TAXONOMIC DISCUSSION. The division of the Oriental fauna of the
subgenus Lutzia into 4 species has long been suspect. Edwards (1932) first
questioned the practice of recognizing 5 species from the Old World, but
Barraud (1934) continued to recognize 4 variable Oriental species. Belkin
(1962) only provisionally recognized the 4 Oriental species of the subgenus
Lutzia, and suggested that they may represent only 1 plastic species or pos-
sibly several distinct sibling allopatric forms. During this study, examina-
tion of over 80 glycerine preparations of Lutzia terminalia from Thailand,
Okinawa, Japan, and the Philippines as well as considerably more adults and
larvae suggests that only 2 variable species of the subgenus Lutzia, fuscanus
and halifaxii, are present in the Oriental region. This conclusion is based
primarily on the anatomy of the male phallosome. Specimens of the typical
halifaxii, raptor, and vorax forms all demonstrate a series of 3 to 10 promi-
nent denticles extending from the middle of the median process (a feature
best observed in lateral aspect). Although color patterns of the scutum, legs,
and abdomen vary considerably, the presence of prominent denticles on the
phallosome are consistent. In contrast, the phallosome of fuscanus does not
exhibit the prominent denticles but instead, possesses striations which form
a suggestion of teeth on the inner margin. Again, color patterns of the
scutum, legs, and abdomen are variable, but the phallosome remains consist-
ent. Glycerine preparations of the male phallosome of the third Old World
species of Lutzia, tigribes Grandpré and Charmoy, were also examined and
found to be distinct from both fuscanus and halifaxii.
BIOLOGY. In general, it appears that larvae of halifaxii inhabit a
broader range of habitats than do those of fuscanus. Belkin (1962) summarized
habitats in the South Pacific and concluded that the larvae are commonly found
Bram: Genus Culex in Thailand 24
in various kinds of temporary and semipermanent ground water habitats and
that they occur frequently in a great variety of artificial containers, as well
as rock pools, stream margins, and tree holes. This species seems to
display a preference for water of high organic content. LaCasse and Yamaguti
(1950) reported that 45 percent of collections of this species in Japan were
from movable artificial containers, whereas Bick (1951) found larvae of this
species most frequently (65 percent of collections) in puddles in New Guinea.
Due to its broad range of habitats, halifaxii is predaceous on a great number
of mosquito species, as well as other arthropods of appropriate size. Host
preferences of the adult females are unknown; however, man has been reported
as an occasional host by several authors (Belkin 1962; LaCasse and Yamaguti
1950; Bonne-Wepster 1954). Most investigators have concluded that although
the larvae are voracious feeders, populations never reach sufficient density
to serve effectively as mosquito control agents. Yasuno (1965) conducted
laboratory studies on vovax (= halifaxii) and concluded that the first stage
larvae have a higher predatory efficiency than those of the second and later
instars. It was also concluded that the settling behavior reduced the chance
of contact between members of the predator species and that cannibalism did
not occur when the prey, which moves more frequently, was present.
SUBGENUS NEOCULEX DYAR 1905
Neoculex Dyar 1905, Proc. ent. Soc. Wash. 7: 48. Type species: Culex
territans Walker.
Maillotia Theobald 1907, Mon. Cul. 4: 274. Type species: Maillotia
pilifera Theobald.
Eumelanomyia Theobald 1909, Colon. Rep. misc. Ser. No. 237: 10. Type
species: Eumelanomyia inconspicua Theobald.
Protomelanoconion Theobald 1910, Mon. Cul. 5: 462. Type species:
Protomelanoconion fusca Theobald.
The adult females cannot be conclusively distinguished from several
closely related subgenera, but, in general, lack acrostichal bristles on the
scutum and exhibit dense scaling on the scutum which is smooth in appearance.
The adult males may be recognized by the characters mentioned above in ad-
dition to the short palpi and the distinctive terminalia. The fourth stage larvae
are recognized by thoracic hair 3-P shorter and slenderer than 1-P and head
hair 14-C single or often dendritic beyond the basal half.
FEMALE. Usually small species, without distinctive ornamentation.
Head. Proboscis unbanded; palpus uniformly dark. Thorax. Dorsocentral
bristles well developed; acrostichal bristles weakly developed, present only
on the extreme anterior edge of the scutum; scales of scutum dense, smooth
in appearance; pleura without dark integumental markings or distinct scale
patches; lower mesSepimeron with or without a bristle; legs all uniformly dark.
Abdomen. Terga with or without narrow, pale, basal bands.
MALE. Similar in general appearance to the female. Head. Palpus
1/2 to 3/4 the length of the proboscis; antenna without specialized scales or
setae. Terminalia. Basimere without scales and without prominent sub-
marginal setae; distimere normal for the genus; phallosome simple, globose
and covered with strong denticles, particularly on the inner margin; proctiger
24 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
crowned with a tuft of short, rather stout spicules; 3 to 5 short, cercal setae
present; basal sternal process not developed.
LARVA. Head. Usually normal in shape and overall chaetotaxy;
hair 1-C strong, tapering to a sharp point; 14-C single, often dendritic beyond
the basal half; 16,17-C either present or absent, when present represented by
extremely minute spicules. Thovax. Hairs 1,2,3-P single, but 3-P about half
the length and width of 1,2-P; integument glabrous. Abdomen. Comb consist-
ing of a broadly triangular patch of fan-shaped scales; siphon variable in length
but always greater than the anal segment, with 5 pairs of subventral tufts;
pecten restricted to the basal third or less of the siphon; saddle completely
ringing segment X; ventral brush consisting of approximately 10 individual
tufts of setae, the basal ones occasionally inserted before the grid.
DISTRIBUTION. The subgenus Neoculex is predominantly Ethiopian,
_ but representatives are known from all major zoogeographical regions. In
Thailand, species of the subgenus, primarily brevipalpis, are distributed
throughout the country.
TAXONOMIC DISCUSSION. Although the classification and systematic
arrangement of species within the subgenus are incompletely understood, it
appears that Neoculex may possibly serve as a unifying link between
Culiciomyia, Lophoceraomyia, and Mochthogenes, with closest affinities to
the latter. The adult females are not only difficult to distinguish from these
3 subgenera, but intrasubgeneric differences are minute. The larva and male
terminalia seem closest to Mochthogenes, but the females are similar to
Culiciomyia in having smooth scaling on the scutum. The 2 species of the
subgenus which occur in Thailand are closely related and will undoubtedly be
assigned to the same group when affinities within the subgenus are finally
deter mined. 3
BIOLOGY. Little is known of the biology of Oriental species of this
subgenus. Larvae are usually found in container habitats such as tree holes,
bamboo internodes, and restricted ground waters. Host preferences of the
females are poorly understood, but man is rarely, if ever, attacked. It is
possible that the females feed to some degree on amphibian and reptilian
hosts. Although there is presently no indication that species of the subgenus
Neoculex play any role in the transmission of arthropod borne viruses in
Thailand, viruses for which neutralizing antibodies have been found in the
human population, have been isolated from species of the subgenus in
Mogambique and South Africa (see for example, Worth and de Meillon 1960).
KEY TO SPECIES OF THE SUBGENUS NEOCULEX IN THAILAND -
ADULT FEMALES
One lower mesSepimeral bristle present; abdominal terga with nar-
row, palesbasal bands . Goss eyo ee Oe. tenuipalpis (p. 29)
Lower mesepimeral bristles absent; abdominal terga completely
Cae osu ee a he he ee brevipalpis (p. 25)
KEY TO SPECIES OF THE SUBGENUS NEOCULEX IN THAILAND -
MALE TERMINALIA
Subapical lobe of the basimere with 2 large and 3 smaller leaf-
Bram: Genus Culex in Thailand 25
sliped eetaes 4.64 Shrew is iss Eales tenuipalpis (p. 29)
Subapical lobe of the basimere with only 1 leaf-shaped seta.......
brevipalpis (p. 25)
KEY TO SPECIES OF THE SUBGENUS NEOCULEX IN THAILAND -
FOURTH STAGE LARVAE
Length of the subventral siphon tufts greater than the width of the
siphon at the point of insertion; individual pecten tooth with the
lateral barbs very numerous and fine on the distal half and
widely spaced and coarse on the proximal half. . tenuipalpis (p. 29)
Length of the subventral siphon tufts less than the width of the
Siphon at the point of insertion; individual pecten tooth with the
lateral barbs all coarse and subequal in size. . .bvevipalpis (p. 25)
CULEX (NEOCULEX) BREVIPALPIS (GILES) 1902
(Figures 6, 7, and 8)
Culex longipes Theobald 1901, Mon. Cul. 2: 68 (2, preoccupied by Fabricius
1850).
Stegomyia brevipalpis Giles 1902, Handb., 2nd. ed. : 384 ('*, 2*).
Culex brevipalpis (Giles): Theobald 1903, Mon. Cul. 3: 146 (¢, 9); Barraud
1924, Indian J. med. Res. 11: 1277 (o*, 2); Barraud 1924, Indian J.
med. Res. 12: 432 (L*).
Culex macropus Blanchard 1905, Les Moustiques: 327 (new name for longipes
Theobald 1901); Barraud 1924, Indian J. med. Res. 11: 1277
(synonymy).
Melanoconion uniformis Leicester 1908, Cul. Malaya: 136 (¢, ?); Edwards
1913, Bull. ent. Res. 4: 237 (synonymy). |
Culex fidelis Dyar 1920, Insec. Inscit. menst. 8: 180 (<); Edwards 1929,
Notul. ent. 9: 4 (Synonymy).
Culex (Neoculex) brevipalpis (Giles): Edwards 1932, in Wytsman, Genera
Insect. , fasc. 194: 194 (taxonomy); Barraud 1934, Fauna Brit. India,
Diptera 5: 348 (o*, 2, L*); Baisas 1935, Philipp. J. Sci. 57: 177 (0%,
2); Lee 1944, Atlas mosq. Larvae Aust. Reg.: 1277 (L*); Bohart
1945, Navmed 580: 72 (o'*, L); Bohart and Ingram 1946, Navmed 1055:
72 (o*, 2, L*); Bonne-Wepster 1954, Roy. trop. Inst. Amst. Spec.
Pub. 111: 109 («*, ¢, L*); Lien 1962, Pacif. Ins. 4: 631 (distribution);
Delfinado 1966, Mem. Amer. ent. Inst. 7: 125 (o*, ?, L*, P*).
The adult female may be recognized by the absence of a lower mesep-
imeral bristle and the absence of narrow pale basal abdominal bands. In the
male the terminalia is distinctive. The fourth stage larva possesses a long
Siphon with the subventral tufts shorter than the width of the siphon at the
point of insertion and the pecten tooth fringed with strong, subequal barbs.
FEMALE. Head, (Figure 6B). Antenna normal, its length approxi-
mately equal to that of the proboscis; palpus slightly darker than the proboscis;
decumbent scales of the vertex dark brown on the occiput, slightly broader
and pale posteriolaterally; erect scales light brown. Thorax. (Figure 6B, D).
26 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig. 6
SSXsc
SNSSS
SSS
SA “|
Sr Vy
i} Gs 4
Mas
Walhe
i
LENG H
i)
Y) Ny
ena
SS
SS
ai
yN J
\\
\ ey
BES
Ss
SS
S. Otawa
C. (Neoculex ) brevipalpis
Bram: Genus Culex in Thailand 27
Integument of the scutum light brown but with faintly darker, longitudinal
stripes in the dorsocentral and acrostichal areas; covered with a uniform
pattern of dark brown scales; integument of the pleuron uniformly light brown
(in Some specimens tinged’with green); distinct scale patches absent but
scattered scales may be present in some areas; lower mesepimeral bristle
absent. Wing. (Figure 6F). Legs. (Figure 6E). All legs uniformly dark
scaled. Abdomen. (Figure 6H). Terga totally dark brown; sterna pale.
MALE. Habitus similar to that of the female. Head. (Figure 6A).
Proboscis with an apparent joint at approximately 2/3 the distance from the
base; antenna plumose; length of the palpus 1/2 to 3/4 the length of the
proboscis, appearing somewhat twisted (figure 6C). Terminalia. (Figure
7C). Subapical lobe of the basimere with 3 strong basal rods which are sub-
equal in length, the proximal 2 hooked apically, the distal 1 gently curved
followed by 3 rather straight setae and a short, narrow leaf; distimere
Slender; lateral plate of the phallosome simple, globose, with from 15 to 25
small denticles principally on the internal margin; proctiger well developed,
crowned with a tuft of strong, broad spines and with 2 or 3 cercal setae
present; basal sternal process not developed.
PUPA. (Figure 7A, B).
LARVA. (Figure 8). Head. Antenna spiculate basad of the insertion
of hair 1-A; 2-6-A inserted at the apex of the shaft; head hair 1-C strong,
robust, tapering to a sharp point; 4-C single, simple, very long; 5, 6-C with
from 2 to 4 branches, pectinate; 7-C with from 5 to 10 branches, pectinate;
14-C single, frequently dendritic on the distal half; 16,17-C absent (as il-
lustrated), or 16-C represented by a minute spicule and 17-C absent. Thovax.
Integument glabrous; hairs 1,2-P single, pectinate, subequal in length; 3-P
single, pectinate, about half the length and width of 1,2-P; 4-P with from 2
to 5 branches, pectinate; 5,6-P single, pectinate; 7, 8-P bifid, pectinate; 14-P
bifid, simple. Abdomen. Integument glabrous; comb consisting of a broad,
triangular patch of from 40 to 60 elongated, fan-shaped scales; siphon index
variable, ranging from 7:1 to 15:1, the siphon very narrow, its sides parallel;
0 pairs of subventral tufts inserted on the siphon beyond the pecten, the apical
tuft somewhat out of line; individual tufts with from 2 to 5 branches, their
length less than the width of the siphon at the point of insertion; pecten
restricted to the basal fifth of the siphon or less, consisting of from 11 to 21
teeth; individual pecten tooth with a pointed apical spine and a series of ap-
proximately 10 coarse, lateral barbs; saddle completely ringing segment X;
ventral brush consisting of 10 individual tufts of setae, all located within the
grid (only 5 tufts illustrated); 2 long and 2 distinctly shorter anal gills present.
Critical examination of several points of chaetotaxy and anatomical
features revealed that the larva of this species is extremely variable. Range
of variation in 96 specimens from 44 collections in Thailand is recorded in
the above description. It is interesting to note the extensive variation in head
Figure 6. C. (Neoculex) brevipalpis. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax; C,
dorsal aspect of the male proboscis and palpus; D, dorsal aspect
a of the female wing; E, anterior surface of the female legs; F,
dorsal aspect of female scutum and scutellum; G, dorsal aspect
of the female abdomen.
28
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C. (Neocu/ex) brevipalpis
14
i
14
Vv
il i
a} 2.) x
bs
. 6. me
312 u
1
10 7
Bram: Genus Culex in Thailand 29
hairs 5,6-C, thoracic hair 4-P, the number of comb scales and pecten teeth,
the number and branching of the siphon tufts, and the siphon index which
averaged 10. 8:1, but ranged from 7:1 to 15:1. Despite this variation, con-
sistent differences have not been found which would suggest an unrecognized
Species complex.
TYPE DATA. Lectotype male of brevipalpis hereby designated: syn-
type male (terminalia mounted and attached to pin), ''Shajahnapur, India, 21
Oct. 1910", in the British Museum. Holotype female of longipes from Singa-
pore in the British Museum. Lectotype male of uwniformis hereby designated:
cotype male (terminalia mounted and attached to pin), ''Kuala Lumpur, Fed.
Malay States, Dr. G. F. Leicester, 1912-35", in the British Museum. Lec-
totype male of fidelis from Los Banos, Philippine Islands in the U. S. National
Museum.
DISTRIBUTION. This species is undoubtedly distributed throughout
THAILAND and specimens have been studied from: Ayutthaya, Chiang Mai,
Chon Buri, Chumphon, Kanchanaburi, Krung Thep, Lampang, Mae Hong Son,
Nakhon Nayok, Nakhon Ratchasima, Narathiwat, Nanthaburi, Phattalung,
Prachuap Khivi Khan, Phare, Ranong, Rayong, Sakhon Nakhon, Satun, Thon
Buri, Trat, Trang, Ubon Ratchatani, and Udon Thani.
In addition to THAILAND, brevipalpis has been reported from the fol-
lowing areas: INDIA, CEYLON, INDOCHINA, CHINA, TAIWAN, RYUKYU-
RETTO, PHILIPPINES, MALAYA, INDONESIA, NEW GUINEA, and the
BISMARCK ARCHIPELAGO. Specimens have been examined in the British
- Museum collection from Kota Belud, NORTH BORNEO**.
During this study the following specimens were examined: 91 males
and 229 females, 97 with their associated larval and pupal skins, and 522
larvae.
BIOLOGY. In Thailand larvae have been collected most frequently
from tree holes and from bamboo stumps and internodes. Additional habitats
included various artificial containers, rock holes, a ditch, anda pond. Baisas
(1935) reported that in the Philippines the larvae are found in forest streams,
but most other authors have reported habitats similar to those found in Thai-
land (see, for example: Bohart andIngram 1946; Bonne-Wepster 1954; and
Lien 1962). Host preferences of the adult females are unknown, but man is
certainly not a normal host; however, Bonne-Wepster (1954) reported that in
New Guinea man may be attacked near the breeding places.
CULEX (NEOCULEX) TENUIPALPIS Barraud 1924
(Figure 9)
Culex tenuipalpis Barraud 1924, Indian J. med. Res. 11: 1278 (*, 2); Bar-
raud 1924, Indian J. med. Res. 12: 433 (L*).
Culex (Neoculex) tenuipalpis Barraud: Edwards 1932, in Wytsman, Genera
Insect. , fasc. 194: 195 (taxonomy); Barraud 1934, Fauna Brit. India,
Diptera 5: 351 (o*, 9, L*); Thurman 1959, Univ. Md. Agric. Exp. Sta.
Bull. A-100: 122 (distribution).
Figure 7. C. (Neoculex) brevipalpis. A,B, dorsoventral view of the pupa;
- C, dorsal aspect of the male terminalia.
30
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
/ rig. o
' a
Ow,
MP
O-L. HWANG
C. (Neoculex) brevipalpis
Bram: Genus Culex in Thailand 31
The adult female is recognized by the presence of 1 lower mesepimeral
bristle and the narrow, pale, basal bands on the abdominal terga. The number
and shape of the setae on the subapical lobe of the basimere of the male
terminalia are diagnostic. The fourth stage larva exhibits very long sub-
ventral tufts on the siphon and possesses distinctive pecten teeth.
FEMALE. Head. Antenna normal, its length approximately equal to
that of the proboscis; proboscis and palpus dark brown; decumbent scales of
the vertex dark brown on the occiput, pale posteriolaterally; erect scales
brown. Thorax. Scutum uniformly covered with dark brown scales; pleuron
uniformly light brown, without distinct scale patches; 1 lower mesepimeral
bristle present. Legs. Anterior surface of the hind femur predominantly
pale, with a narrow apical dark band which extends proximally for a short
distance along the dorsal margin; hind tibia and tarsus, as well as the fore
and mid legs uniformly dark. Abdomen. Terga dark brown, with narrow,
pale basal bands; sterna brown, somewhat lighter basally.
MALE. Head. Proboscis with an apparent joint at approximately 2/3
the distance from the base; length of the palpus slightly greater than half the
length of the proboscis, not noticeably twisted as in brevipalpis. Terminalia.
(Figure 9C). Subapical lobe of the basimere prominent, with 3 strong, sub-
equal, hooked basal rods, a very slender leaf, 2 rather broad leaflets with
fringed posterior margins, and 3 rather large, symmetrical, subequal leaf-
lets; distimere slender, normal; lateral plate of the phallosome simple,
globose, with approximately 20 strong denticles, particularly on the internal
- margin.
LARVA. Specimens not available for study; figure 9A,B is after Bar-
raud (1924b).
TYPE DATA. Lectotype male of tenuipalpis hereby designated: syn-
type male (terminalia slide mounted), 'India: Sureil, E. Himalayas, 5,500
ft., X 1922, P. J. Barraud, BM 1923-585" in the British Museum.
DISTRIBUTION. In THAILAND, tenuipalpis is known from Ngao,
Lampang and Doi Suthep, Chiang Mai. In addition to the type locality in
INDIA, this species has also been recorded by Brug (1931) from 1 adult fe-
male and 3 larvae collected on JAVA. During this study 7 male specimens
were examined in addition to the lectotype male.
TAXONOMIC DISCUSSION. The male terminalia of this species is
similar to that of hayashii Yamada, but slight differences exist in the shape
of the setae on the subapical lobe of the basimere; iayashii also lacks the
pale, basal tergal bands, but from the published description, the larvae ap-
pear to be very similar.
BIOLOGY. Barraud (1934) recorded the larval habitat of tenuipalpis
as roadside pools. The specimens which have been examined from Thailand
have been collected as adults, either resting or with a net during February,
March, and June. Feeding preferences and habits of the adult females are
unknown.
Figure 8. C. (Neoculex) brevipalpis. Fourth stage larva: dorsoventral
view of the head and prothorax, and lateral aspect of the
< terminal abdominal segments.
32 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
BE
nae: asssssasnsse ) =
SSS3 3) RSS
SNS 4 SS
TNS sie2, 7
LP EAD Z Carn ip e;
SSG CSM POPLIN tho re
So Fey) ee
. (Neoculex) tenuipalpis
Bram: Genus Culex in Thailand 33
SUBGENUS MOCHTHOGENES Edwards 1930
Culex (Mochthogenes) Edwards 1930, Bull. ent. Res. 21: 305. Type species:
Culex malayi Leicester.
The adult females exhibit strongly developed acrostichal bristles, one
strong, lower mesepimeral bristle, and lack banding patterns on the pro-
boscis, legs, and abdominal terga. The adult males may be recognized by
the characters mentioned above as well as the palpi which are less than 1/4
the length of the proboscis. The fourth stage larvae possess a strong siphon
with well developed pecten which bears distinctive teeth, head hair 1-C well
developed, and thoracic hair 3-P more slender than 1-P and less than half its
length; head hairs 5,6-C are short and weak and 14-C is bifid.
FEMALE. Very small species devoid of distinctive ornamentation.
Head. Proboscis and palpus completely dark; antenna longer than the pro-
boscis. Thorax. Acrostichal, anterior dorsocentral, posterior dorsocentral,
supraalar, and prescutellar bristles all well developed; pleuron without distinct
scale patches; 1 strong, lower mesepimeral bristle present; all legs completely
dark.
MALE. Head. Palpus less than 1/4 the length of the proboscis; anten-
na plumose, without specialized scales or setae. Terminalia. Basimere
without scales or distinctive submarginal setae; subapical lobe having strongly
developed setae; distimere either bifid or normal; phallosome simple, elongate,
with or without denticles; proctiger crowned with a tuft of rather strong, short
spines; 2 or 3 short cercal setae present; basal sternal process not developed.
LARVA. Head. Head hair 1-C well developed, tapering gradually to
a sharp point; 5,6-C short, weak, 5-C about half the length of 6-C; 14-C
bifid, the branches strongly divergent; 16,17-C present, represented by
minute spicules. Thorax. Hairs 1,2,3-P single, 3-P approximately half the
length and width of 1,2-P. Abdomen. Comb consisting of a broadly triangu-
lar patch of scales; pecten restricted to the basal fourth of the siphon; in-
dividual pecten tooth with several broad basal barbs and a distal lateral series
of fine, parallel spines; saddle completely ringing segment X; ventral brush
consisting of 10 or more tufts.
DISTRIBUTION. The subgenus Mochthogenes is restricted to the Old
World tropics. Species of the subgenus are found throughout Thailand.
TAXONOMIC DISCUSSION. Mochthogenes demonstrates closest af-
finity to the subgenera Neoculex and Lophoceraomyia. The short palpus of
the male is quite distinctive and species in Thailand possess a unique pecten
tooth in the fourth stage larva. The subapical lobe of the basimere in the
male terminalia usually exhibits extensively developed processes and, at this
stage of our knowledge, the male terminalia presents the only completely
reliable structures for species diagnosis within the subgenus in Southeast
Asia.
BIOLOGY. Bionomics of the adults of this subgenus are virtually un-
known; however, females have never been reported feeding on man. Larvae
Figure 9. C. (Neoculex) tenuipalpis. A,B, dorsal view of the fourth stage
~ larva (after Barraud 1924b); C, dorsal aspect of the male
‘elit terminalia.
34 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
have been collected from a wide variety of habitats in Thailand, ranging from
stream margins to rock pools, wells, and ponds.
KEY TO SPECIES OF THE SUBGENUS MOCHTHOGENES IN THAILAND -
ADULT FEMALES!
Decumbent scales of the vertex uniformly dark; anterior surface of -
the hind femur pale; abdominal sterna dark...... foliatus (p. 34)
Decumbent scales of the vertex uniformly pale; anterior surface of
the hind femur dark; abdominal sterna mainly pale. .malayi (p. 39)
KEY TO SPECIES OF THE SUBGENUS MOCHTHOGENES IN THAILAND -
MALE TERMINALIA :
1, Distimere furcate........ a es wactiaied elie sows malayi (p. 39)
Ere RUIN Gi Sais ES WR WTO WR OE Se Bee oe 2
2 Lateral plate of the phallosome with distinct denticles, particularly
on the inner surface; subapical lobe of the basimere with 4 leaf-
shaped setae present... 66 seek ee ees . foliatus (p. 34)
Lateral plate of the phallosome without distinct denticles; sub-
apical lobe of the basimere without a leaf-shaped seta.........
hinglungensis (p. 377)
KEY TO SPECIES OF THE SUBGENUS MOCHTHOGENES IN THAILAND -
FOURTH STAGE LARVAE!
Comb possessing 2 distinct types of scales, some fan-shaped and
others with the median apical spine broader and longer than
the other spines; head hairs 5,6-C simple....... malayi (p. 39)
All comb scales fan-shaped, fringed with subequal spines; head
hairs 6 GO peclinaten. 6.65 ce ee ads, Ge foliatus (p. 34)
CULEX (MOCHTHOGENES) FOLIATUS BRUG 1932
(Figures 10 & 12A)
Culex (Culex) aot th var. foliatus Brug 1932, Bull. ent. Res. 23: 82 (o*,
L*, P*).
Culex (Mochthogenes) chiyutoi Baisas 1935, Philipp. J. Sci. 57: 176 (o*, 9);
Bohart 1945, Navmed 580: 73 (o‘*); Delfinado 1966, Mem. Amer. ent.
Inst. 7: 129 (**). NEW SYNONYMY.
Culex (Mochthogenes) castrensis var. foliatus Brug: Peters and Dewar 1956,
Indian J. Malar. 10: 46 (o'*).
Culex (Mochthogenes) foliatus Brug: Bohart 1946, Navmed 961: 16 (distribu-
1
The adult female and larva of hinglungensis are unknown.
Bram: Genus Culex in Thailand 35
tion, key).
The adult female may be distinguished from the other members of the
subgenus in Thailand by the uniformly dark brown decumbent scales of the
vertex, the pale scaled hind femur, and the dark abdominal sterna. The
larva is distinctive in that all comb scales are fan-shaped and fringed with
subequal spines. The male terminalia is characterized by the distinctive
chaetotaxy of the subapical lobe of the basimere. :
FEMALE. A very small species without prominent ornamentation or
color patterns. Head. Decumbent scales of the vertex uniformly dark brown,
but with some lighter scales on the posterior orbital line; erect scales uni-
formly dark brown. Thorax. Integument of scutum and scutellum uniformly
dark brown but with faintly lighter longitudinal stripes in the dorsocentral
area; covered with a uniform pattern of bronze-brown scales; integument of
the pleuron uniformly dark brown, in some specimens distinctly tinged with
ereen, particularly in the areas of the meron, precoxa, and lower meSepimeron;
scales absent; a series of approximately 5 upper mesepimeral bristles present,
several upper sternopleural bristles, 1 posterior sternopleural bristle, and 1
strong lower mesSepimeral bristle present; except for the pale scaled anterior
surface of the hind femur, all legs uniformly dark scaled. Abdomen. Terga
and sterna totally dark. ,
MALE. Generally marked as the female. Head. Palpus less than
1/5 the length of the proboscis, completely dark. Terminalia. (Figure 12A).
Subapical lobe of the basimere rather flattened with 3 strong rods present,
the basal 1 expanded medially and strongly elbowed with a hooked apex, median
rod bent with an apical hook, distal rod gently curved and hooked apically;
following the rods is a rather broad, blunt seta and 4 subequal, finely striated,
rather symmetrical leaves; distimere swollen, enlarged distally, witha
variable number of randomly placed setae; lateral plate of the phallosome
simple, elongated, with small denticles, particularly at the apex and on the
internal margins.
LARVA. (Figure 10). Head, Antenna spiculate basally and slightly
beyond insertion of hair 1-A; antennal shaft constricted beyond insertion of
1-A; hairs 2-6-A normal for the genus, but 2, 3-A inserted slightly below the
apex of the antenna; head hair 1-C robust, darkly pigmented, tapering to a
sharp point; 4-C with from 2 to 4 branches, simple; 5-C bifid, finely pecti-
nate, very short and slender; 6-C bifid, pectinate, robust; 16,17-C repre-
sented by minute spicules. Thorax. Integument glabrous; hairs 1,2-P
single, pectinate, subequal in length; 3-P single, simple, about half the length
and width of 1,2-P; 4-P small, with 4 or 5 branches, simple; 5, 6-P single,
pectinate; 7-P single or bifid, pectinate; 8-P single, pectinate; 14-P bifid,
simple. Abdomen. Integument glabrous; comb consisting of a triangular
patch of approximately 35 somewhat elongate, fan-shaped scales which are
fringed with subequal spines; siphon index approximately 6:1, the sides of
the siphon gradually tapering to a truncate apex; 6 pairs of subventral tufts
evenly spaced on the siphon beyond the pecten; individual tufts 3 or 4
branched, their lengths exceeding the width of the siphon at the point of
insertion; pecten restricted to the basal fourth of the siphon, consisting of
approximately 8 to 10 teeth; individual pecten tooth rather elongated,
fringed on the distal half with a series of 10 or more fine spines and with 3 or
4 rather coarse barbs on the proximal half; saddle completely ringing seg-
[
36 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.10
WW)
HY)
V7
NY
i)
ay
cs
[*)
,
NN Ni if
: Sv a
VS A
NG Lp
_
— Li
—G a
w& : :
“JTL. HWANG
C.(Mochthogenes ) foliatus
Bram: Genus Culex in Thailand 3a
ment X.
TYPE DATA. The holotype male of foliatus (terminalia mounted and
attached to pin) from Bandoeng, Java is in the British Museum. The original
description of chiyutoi designated one male type, one female type, and five
male cotypes from Kolambugan, Lanao Province, Mindanao, Philippines. Un-
fortunately, all specimens in the type series have been either lost or destroyed
except for a single male which is undoubtedly one of the original 5 cotype
males. Therefore, a lectotype is hereby designated: male cotype (terminalia
slide mounted), "R 33, U.S.N.M. paratype no. 51392, slide no. 44. VII. 22b",
in the U. S. National Museum.
DISTRIBUTION. In THAILAND, /foliatus has been collected from:
Chiang Mai, Chon Buri, Chumphon, Mae Hong Son, Nakhon Nayok, Nakhon
Ratchasima, Narathiwat, Prachuap Khiri Khan, Phare, and Songkhla. This
species has also been recorded from JAVA, CHINA, NEPAL, and the PHILIP-
PINES (as chiyutoi). Specimens have been examined in the British Museum
collection from Kota Belud, NORTH BORNEO**.
During this study the following specimens have been examined from
Thailand: 29 males and 32 females (32 with their associated larval and pupal
skins), and 119 larvae.
TAXONOMIC DISCUSSION. Brug (1932) originally described foliatus
as a variety of castrensis. Bohart (1946) treated foliatus as a valid species,
but subsequent authors (Peters and Dewar 1956; Stone e¢ al. 1959) continued
to recognize foliatus as either a variety or subspecies of castrensis. Com-
parison of the 2 holotype males has revealed that in light of modern taxonomic
practice, foliatus and castrensis should be recognized as distinct and valid
species.
Examination of the holotype male terminalia of folizatus and the lecto-
type male terminalia of chiyutoi has revealed these forms to be conspecific.
The distinctive setae on the subapical lobe of the basimere are identical in
both forms, and the larva of chiyutoi (first described by Delfinado 1966) con-
forms to associated larval skins of foliatus from Thailand.
BIOLOGY. Larvae of this species have been reported exclusively
from pools, generally with clear water (Brug 1932; Feng 1938). Although
most collections of foliatus in Thailand were from stream pools, rock pools,
and puddles, collections were also made from elephant hoof prints, seepage
springs, a bamboo internode, anda bamboo pot. In the Philippines larvae
have been reported from tree holes and small ground pools (as chiyutoi).
Host preferences of the adult females are unknown, but they have not been
reported to feed on man.
CULEX (MOCHTHOGENES) HINGLUNGENSIS CHU 1957
(Figure 12B)
Culex (Mochthogenes) hinglungensis Chu 1957, Acta. zool. Sinica 9:159 (c’);
Chu 1958, Indian J. Malar. 12: 111 (¢*).
Figure 10. C. (Mochthogenes) foliatus. Fourth stage larva: dorsoventral
aspect of the head and prothorax, and lateral aspect of the
< terminal abdominal segments.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
38
Fig. 11
C. (Mochthogenes) malayi
Bram: Genus Culex in Thailand 39
The male terminalia of this species is characterized by the absence of
denticles on the lateral plate of the phallosome and by the number and arrange-
ment of setae on the subapical lobe of the basimere.
FEMALE. Unknown.
MALE (After Chu 1958). A small dark mosquito, closely related to C.
(Mochthogenes) pluvialis Barraud, but differing in the male terminalia. Head.
Antenna, palpus, and proboscis dark brown, the palpus about 1/6 the length of
the proboscis; a large area of flat scales with some narrow, curved and up-
right scales on the vertex and occiput. Thorax. Scutum and scutellum covered
with narrow, curved, dark brown scales; pleuron dark brown, without scales;
wings dark scaled; legs dark brown. Abdomen. Terga dark brown, sterna
paler. Terminalia. (Figure 12B). Subapical lobe of the basimere weakly
developed, with a reduced number of setae; lateral plate of the phallosome
simple, rather short, without distinctive denticles.
LARVA. Unknown.
TYPE DATA. The holotype male from Hing-lung, Hainan Island,
China is in the Department of Parasitology, Second Military Medical Univer-
sity, (precise location not given) China.
DISTRIBUTION. Apart from the type locality a single male has been
collected in a light trap at Chiang Dao, Chiang Mai, THAILAND**.
TAXONOMIC DISCUSSION. Due to the abbreviated original and sub- ©
sequent descriptions and the unavailability of the holotype for comparison,
hinglungensis is only tentatively recognized from Thailand, pending exami-
nation of a more complete series. As far as is known, the species is
represented only by the holotype male and the single male from Thailand.
BIOLOGY. The holotype male was collected in the crevice of rocks
near mountain streams on Hainan Island during May. The male specimen
from Thailand was collected at a light trap next to a stream during April.
CULEX (MOCHTHOGENES) MALAYI (LEICESTER) 1908
(Figures 11 & 12C)
Aedes malayi Leicester 1908, Cul. Malaya: 184 (¢, 9).
Aedes nigrescens Theobald 1907, Mon. Cul. 4: 540 (preoccupied by Theobald
1907: 248).
Aioretomyia aedes Leicester 1908, Cul. Malaya: 189; Edwards 1917, Bull.
ent. Res. 17: 228 (synonymy).
Micraedes malayi (Leicester): Edwards 1917, Bull. ent. Res. 17: 228
(taxonomy); Edwards 1922, Bull. ent. Res. 13: 92 (taxonomy).
Culex malayi (Leicester): Barraud 1924, Indian J. med. Res. 11: 1282 (*);
Borel 1926, Arch. Insts. Pasteur Indo-Chine 3-4; 43 (o*, 9, L*);
Borel 1930, Monogr. Coll. Soc. Path. exot. 3: 368 (o*, 2, L*).
Culex (Mochthogenes) malayi (Leicester): Edwards 1930, Bull. ent. Res. 21:
305 (taxonomy); Barraud 1934, Fauna Brit. India, Diptera 5: 358 (0'*,
°,L); Li and Wu 1935, Yearb. Bur. Ent. Chekiang 4: 98 (L*); Brug
Figure 11. C.(Mochthogenes) malayi. Fourth stage larva: dorsoventral
aspect of the head and prothorax, and lateral aspect of the
terminal abdominal segments.
40 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
ia 2.
C.(Mochthogenes) foliatus
C. (Mochthogenes ) malayi
Bram: Genus Culex in Thailand 41
1939, Tijdschr. Ent. 82: 111 (L*); Lee 1944, Atlas Mosq. Larvae
Aust. Reg. 90 (L*); Galliard and Ngu 1949, Ann. Parasit. num.
comp. 24: 502 (o*, L*); Bonne-Wepster 1954, Roy. trop. Inst. Amst.
Spec. Pub. 111: 111 (¢,?, L*); Hsieh and Liao 1956, Acta ent. Sinica
6: 376 (L*); Lien 1962, Pacif. Ins. 4: 631 (distribution).
The adult female can be distinguished by the pale decumbent scales of
the vertex, the dark hind femur, and the pale sterna. The fourth stage larva
possesses 2 types of scales in the comb and hair 4-P is dendritic. The male
terminalia is easily recognized by the furcate distimere and the chaetotaxy of
the subapical lobe of the basimere.
FEMALE. A small species without prominent ornamentation or color
patterns. Head. Proboscis and palpus uniformly dark brown; decumbent
scales of the vertex pale; erect scales dark brown. Thorax. Scutum uniform-
ly dark brown with faintly paler longitudinal stripes on the dorsocentral area,
and clothed with a uniform pattern of bronze-brown scales; integument of the
pleuron uniformly dark brown, but frequently tinged with green in the areas of
the meron, precoxa, propleuron, and lower mesSepimeron; scale patches
absent; 7 to 10 weak upper mesepimeral bristles and 1 strong lower mesep-
imeral bristle present. Legs. All legs uniformly light to dark brown without
indication of pale bands. Abdomen. Terga dark brown; sterna I-IV with a
mixture of pale and dark scales, sterna V-VII completely pale.
MALE. Habitus generally as in the female. Head. Length of the
palpus 1/5 or less than that of the proboscis. Terminalia. (Figure 12C).
Subapical lobe of the basimere flattened, with strongly developed setae; with
3 subequal, hooked basal rods followed by a very broad, strong, striated, and ©
characteristically shaped leaf, and 2 clavate setae, 1 of which is distinctly
serrate; distimere characteristically furcate; lateral plate of the phallosome
simple, elongate, covered with small, but distinct, denticles particularly at
the apex and on the internal margin.
LARVA. (Figure 11). Head. Antenna spiculate basally and slightly
beyond insertion of hair 1-A; antennal shaft constricted beyond insertion of
1-A; 2,3-A inserted somewhat below the apex of the shaft; head hair 1-C robust,
darkly pigmented, tapering to a sharp point; 4-C bifid, simple; 5-C with from
2 to 4 branches, simple, short and very fine; 6-C with from 2 to 4 branches,
simple, longer than 5-C; 7-C with 9 branches, pectinate; 16,17-C represented
by extremely minute spicules. Thorax. Integument glabrous; hairs 1,2-P
Single, finely pectinate, subequal in length; 3-P single, finely pectinate, about
half as long and wide as 1,2-P; 4-P with from 4 to 6 very fine, pectinate
branches; 5,6-P single, pectinate; 7-P bifid, pectinate; 8-P single, pectinate;
14-P bifid, simple. Abdomen. Integument glabrous; comb consisting of a
triangular patch of approximately 20 scales; posterior scales with a promi-
nent apical spine and fringed with minute lateral spicules, anterior scales
fan-shaped, fringed with subequal spines; siphon index approximately 6:1, the
sides gradually tapering to a truncate apex; 6 or 7 pairs of subventral tufts
inserted beyond the pecten; individual tufts with 7 or more branches, their
Figure 12. Dorsal aspect of male terminalia: A, C. (Mochthogenes)
< foliatus; B, C. (Mochthogenes) hinglungensis; C, C.
(Mochthogenes) malayi.
42 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
length exceeding the width of the siphon at the point of insertion; pecten
restricted to the basal fourth of the siphon, consisting of approximately 9
teeth; individual pecten tooth rather long and slightly curved, fringed with
long, narrow, Sharply pointed and numerous barbs on the distal half and
short, robust barbs on the basal half; saddle completely ringing segment X;
ventral brush consisting of from 12 to 14 tufts usually inserted within the
erid.
TYPE DATA. Lectotype male of malayi hereby designated: cotype
male (terminalia slide mounted), "Kuala Lumpur, Fed. Malay States, 29/9/
03, G. F. Leicester, clear pool, circular road", in the British Museum.
Holotype male of aedes (terminalia mounted and attached to pin) from Kuala
Lumpur in the British Museum. Holotype male of nigrescens (terminalia
slide mounted) from Castle Rock, Bombay, India in the British Museum.
DISTRIBUTION. In THAILAND malayi has been collected from:
Chiang Mai, Chon Buri, Krung Thep, Lampang, Nakhon Ratchasima, Phet
Buri, Phare, Satun, and Songkhla. This species has also been recorded
from: MALAYA, INDIA, CEYLON, NEPAL, MALDIVE ISLANDS, ANDAMAN
ISLANDS, BURMA, INDONESIA, TAIWAN, and NEW GUINEA. Specimens
have been examined in the British Museum collection from Kota Belud, NORTH
BORNEO**. Delfinado (1966) has cast doubt on the validity of records from
the PHILIPPINES.
During this study the following specimens have been examined from
Thailand: 68 males and 19 females (34 with associated larval and pupal
skins) and 59 larvae.
TAXONOMIC DISCUSSION. On the basis of the male terminalia,
malayi exhibits its closest affinity to yeageri Baisas and laureli Baisas both
of which are endemic to the Philippine Islands; however, differences in the
shape of the setae on the subapical lobe of the basimere are diagnostic.
BIOLOGY. Larvae have been reported from stream pools, ditches,
swamps, wells, and similar ground water habitats (Carter and Wijesundara
1948; Bonne-Wepster 1954; Lien 1962). In Thailand specimens have been
collected most frequently from stream margins, stream pools, and from
Slowly moving water in ditches. Host preferences of the adult females are
unknown, but they have never been reported to attack man.
SUBGENUS LOPHOCERAOMYIA THEOBALD 1905
Lophoceraomyia Theobald 1905, J. Bombay nat. Hist. Soc. 16: 245 (April).
Type species: Lophocevaomyia uniformis Theobald.
Lophoceratomyia Theobald 1905, Ann. hist.-nat. Mus. hung. 3: 93 (June,
lapsus),.
Philodendromyia Theobald 1907, Mon. Cul. 4: 623. Type species:
Philodendromyia barkerii Theobald.
Cyathomyia Meijere 1910, Ann. bot. Gdn. Buitenz., Suppl. 3: 921. Type
species: Cyathomyia jenseni Meijere.
The adult females are most difficult to characterize and to separate
from the subgenera Mochthogenes, Neoculex, and Culiciomyia; proboscis un-
banded, pleuron without distinct scale patches, acrostichal bristles weakly
developed except at extreme anterior margin of the scutum, scales of the
Bram: Genus Culex in Thailand 43
scutum narrow, sparse, the pattern rather rough in appearance. The adult
males are immediately recognized by the presence of distinctly modified
scales and setae on flagellomeres V-IX or X. The fourth stage larvae possess
a well developed pecten; ventral brush consisting of 10 or more tufts; thora-
cic hair 3-P slenderer than 1-P and approximately half its length; head hair
14-C bifid, its branches strongly divergent; head hairs 5,6-C strong, long,
subequal in length.
FEMALE. Usually small to moderately sized species. Head. Pro-
boscis and palpus completely dark; length of antenna slightly greater than the
length of the proboscis; decumbent scales of the vertex usually dark on the
occiput and becoming slightly paler at the orbital line; erect scales usually
uniformly dark. Thorax. Scutum sparsely covered with narrow dark scales;
acrostichal bristles usually absent, weakly developed at the anterior margin
of the scutum; dorsocentral bristles well developed; pleuron usually without
distinct scale patches (never as extensive as seen in the subgenus Culex);
lower mesepimeral bristle present or absent. Wing. All dorsal wing scales
dark, sparse except towards the apex. Legs. Tibiae and tarsi of all legs
uniformly dark scaled; anterior surface of the hind femur often predominantly
pale, with an apical dark band which extends proximally as a narrow dark
stripe along the dorsal border. Abdomen. Terga usually uniformly dark
scaled, occasionally with narrow, basal pale bands; sterna variable.
MALE. Generally marked as the female. Head. Length of the palpus
ereater or less than that of the proboscis; antenna usually with modified
scales and setae on flagellomeres V-IX or X (always present on flagellomeres
VII and VIII). Terminalia. Basimere with strongly developed submarginal
setae, usually in a row; subapical lobe with prominent and characteristic
setae, at least 1 of which is usually leaf-shaped; distimere well developed, its
shape variable although usually slender and with 2 small setae inserted on the
distal third; phallosome consisting of 1 or 2 lateral plates; proctiger well
developed, crowned with a tuft of several rather strong, short bristles; usually
2 or 3 cercal setae present; basal sternal process not developed.
LARVA. Head. Integument moderately pigmented, the antennal shaft
usually concolorous with the head capsule, but with a variable, narrow, dark
basal band; hair 1-C darkly pigmented, tapering to a sharp point, its length
usually equal to approximately half the distance between the bases of the pair;
principle head hairs well developed; 5, 6-C strong, long, subequal in length;
14-C bifid, its branches strongly divergent; 16,17-C present or absent, when
present represented by minute spicules. Thorax. Integument with or without
spicules; hairs 1,2,3-P usually single, 3-P very fine, considerably shorter
than 1,2-P, occasionally bifid. Abdomen. Comb well developed, usually with
fan-shaped scales (except in minor in which the scales of the posterior row
are sharply pointed); siphon well developed, with 3 or 4 pairs of subventral
tufts usually inserted in a line; pecten always present, restricted to the basal
third of the siphon or less (except in curtipalpis in which the pecten covers
the basal half of the siphon); ventral brush consisting of 10 or more tufts; sad-
dle completely ringing segment X.
DISTRIBUTION. The subgenus Lophocevaomyia is restricted pre-
dominantly to the Oriental, Indomalayan, and Australasian regions of the Old
World. In Thailand, representatives of the subgenus are found throughout
the country. Colless (1965) has postulated that the subgenus originated in
Southeast Asia as a derivative of the stem that includes Neoculex, Moch-
44 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
thogenes, and Culiciomyia.
TAXONOMIC DISCUSSION. As indicated above, the subgenus Lopho-
cevaomyia demonstrates its closest affinity to Neoculex, Mochthogenes, and
Culiciomyia, as well as the monotypic subgenus Thaiomyia. The adult fe-
males of these 5 subgenera are very Similar, and the male terminalia, larvae,
and pupae all show close relationships. Future investigations in Southeast
Asia and other regions where these subgenera are found may well indicate the
necessity for a realignment of species within a modified subgeneric frame-
work.
Species within the subgenus Lophocevaomyia have been divided into the
2 major groups proposed by Colless (1965) as opposed to the 3 groups rec-
ognized by Barraud (1934). This classification, although somewhat complex,
seems to clearly define the anatomical and biological differences of the 2
major groups, while recognizing the supposed phylogenetic affinities within
the mammilifer group.
BIOLOGY. Larvae of species belonging to the brevipalpus subgroup
of the mammilifer group are found predominantly in pitcher plants (Nepenthes
spp.). Species of the fraudatrix group may be found as larvae primarily in
various ground water habitats; larvae of the mammilifer subgroup are most
frequently found in natural container habitats (e.g. tree holes, bamboo inter-
nodes, rock holes, etc.), but are not uncommon in ground water habitats as
well. Very little is known of the biting habits of the adult females. For the
most part, they probably prefer avian hosts and man rarely, if ever, serves
as a host for any species within the subgenus. Three arthropod-borne
viruses (Bakau, Ketapang, and Bebaru viruses) have been isolated from
species of the subgenus Lophoceraomyia in Malaya, but the mosquito species
were not determined, nor were host relationships fully investigated; however,
antibodies to these viruses have been identified from humans in Malaya
(Anonymous 1960; in this report Bakau virus is designated MM-2325, Keta-
pang is MM-2549, and Bebaru is MM-2354).
KEY TO SPECIES OF THE SUBGENUS LOPHOCERAOMYIA IN THAILAND -
ADULT MALES
Ls Phallosome with both an internal process and a spinose or toothed
dorsal process; torus of antenna asymmetrical or witha
distinct prominence or protuberance on the inner surface;
palpus without finger-like basal processes. . (mammilifer group) 2
Phallosome with a pointed, toothless dorsal process only; torus of
antenna not asymmetrical and without a distinct prominence;
palpus with finger-like basal processes. .... (fraudatrix group) 16
2(1). Apex of distimere noticeably expanded; basimere without long,
prominent submarginal setae......... (brevipalpus subgroup) 3
Apex of distimere normal; basimere with long, prominent sub-
Mmareinal Seta oe ee ee Pe eS (mammilifer subgroup) 4
3(2). Flagellomere V with a tuft of 2 or 3 dark brown, very long, fila-
mentous scales, followed laterally by 3 or 4 shorter, narrow,
pointed scales; palpus, at most, half the length of the pro-
Oe. Ge Re ia ae eee curtipalpis (p. 76)
4(2).
5(4).
6(5).
15).
8(7).
9(8).
10(9).
11(10).
Bram: Genus Culex in Thailand 45
Flagellomere V with a tuft of 6 or 7 golden, shorter and broader
tapering scales; palpus from 1/2 to 3/4 the length of the pro-
WON ee el ee ee ae Oe lucaris (p. 79)
Dorsal process of the phallosome very broad, the distal half
covered with numerous strong teeth; internal process very small
and toothed; torus of antenna without a mammiliform protuber-
ance, Slightly swollen on inner surface, and with a circum-
ferential, almost slit-like depression........ wilfredi (p. 110)
Dorsal process of the phallosome narrower, spinose or spiculate
at apex and with lateral denticles on the inner surface; internal
process curved and pointed; torus of antenna with a distinct
prominence or protuberance on the inner surface and without
A Blitwie Ger eset re oe ere ee ce fw 6 5)
Flagellomere V with a conspicuous tuft having from 1 to 4 long,
DIOR Sete a ee ee ee ee ee ay 6
Basimere with from 6 to 8 long, prominent submarginal setae; in-
ternal process of the phallosome projecting beyond the apex of
the dorval process. 6. ee mammilifer (p. 98)
Basimere with 3 long, prominent submarginal setae; internal
process not projecting beyond the apex of the dorsal process... .
bholeter (p. 103)
Basimere with a single row of long prominent, submarginal setae. . .8
Basimere with 2 irregular rows of long, prominent, submarginal
SO Ee re ee ee ae ee ee ee a te 15
BOAO Pe OS ee Oe CE ON OE TON ho at ek ie 9
Flagellomere IX without a tuft of obviously modified setae or with,
at most, a few setae shorter than the rest, but not stronger or
OTREPW Ibe MOCIOG og i es ok a ee ee 13
Flagellomere VII with an internal tuft of from 3 to 5 gently sigmoid,
specialized setae eI PO Ea Se Oa eukrines (p. 85)
Flagellomere VII with an internal tuft of 5 or more strongly
sigmoid or kinked, Specialized setae’. Fe es es 10
Lower mesepimeral bristle absent; flagellomere V with a tuft of
from 1 to 4 very long, filamentous scales, their apices reaching
beyond the tuft on flagellomere VI........ bengalensis (p. 82)
Lower mesepimeral bristle present; flagellomere V not as above. . .11
Flagellomere V with a tuft of long, yellowish hairs; distimere with
1 or 2 setae on the proximal third; subapical lobe of the basimere
with the internal rod slightly expanded subapically, followed by
arather broad, gently curved, slightly sclerotized apex.......
peytoni (p. 100)
46
12(11).
13(8).
14(13).
15(7).
16(1).
17(16).
18(17).
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Flagellomere V with a tuft of short, narrow, seta-like scales; disti-
mere without setae on the proximal third; subapical lobe of the
basimere with the internal rod slightly expanded apically, then
narrowing to a filamentous tip, often with a fine apical hook. ... 12
Basal half of proboscis, distad of the joint, with a slightly swollen
portion bearing numerous, long, curved hair-like, ventro-
TOLOrS) BOlIG 6kcs khaled minor form bicornutus (p. 99)
Proboscis not swollen and without ventrolateral setae........... '
minor (p. 96)
Flagellomere V with a tuft of long, narrow, tapering, pointed
scales, their apices reaching almost to the tuft on flagellomere
Vill; flagellomere VIII with an internal tuft of strongly curved,
specialized SctIae . we ee ew ganapathi (p. 90)
Flagellomere V with a tuft of short, very narrow, acute scales,
their apices, at most, reaching to the tuft on flagellomere VI;
flagellomere VIII with an internal tuft of rather straight or
gently curved, specialized setae. ....... 20.2000 cccccvcs 14
Flagellomere VIII with an internal tuft of long, gently curved,
specialized setae; internal process of the phallosome projecting
beyond the apex of the dorsal process.......... traubi (p. 108)
Flagellomere VII with an internal tuft of long, rather straight
specialized setae; internal process of the phallosome not
projecting beyond the apex of the dorsal process.............
spiculosus (p. 105)
Internal process of the phallosome projecting beyond the apex of
the dorsal Or Ocess. o.oo es fuscosiphonis (p. 87)
Internal process of the phallosome not projecting beyond the apex
Gr CHE MORSGr PTOCENS . be ee ee incomptus (p. 81)
Basimere with a patch of prominent submarginal setae...........
cinctellus (p. 55)
Basimere with the prominent submarginal setae inaline........ 17
Dorsal process of the phallosome with reticular markings on the
entire surface; submarginal setae fine; flagellomeres V and VI
without tufts of scales or specialized setae... . infantulus (p. 58)
Dorsal process of the phallosome smooth; submarginal setae
strong; flagellomeres V and VI each with a tuft of scales or
Specialized Sctaew bic bavi AO ai ie & eS eGhet ks 18
Flagellomere V with a rather inconspicuous tuft of narrow, tapering,
pointed scales; lower mesepimeral bristle absent; integument
of the scutum often with a reddish tint...... rubithoracis (p. 69)
Flagellomere V with a conspicuous tuft of broad scales; lower
mesepimeral bristle present; integument of the scutum without
BOO CIR es A OR Oe ee 19
Bram: Genus Culex in Thailand 47
19(18). Flagellomere V with the more ventral scales rather long and ex-
panded apically; abdominal sternum VII with white scales,
markedly contrasting with the general dark integument and
scaline® on the other BtOrWA .5culk 6 oe a ete eos alphus (p. 53)
Tuft of scales on flagellomere V and abdominal scaling other-
WIRO Ly aie od ook ko eee be © ee Gi ee dae gs! fae cae 20
20(19). Flagellomere V with 8 or more dorsal scales which are subequal
Wid DULY COO 6s es See ce ere ie es te A ee mw 21
Flagellomere V with 6 or less dorsal scales which vary in length
and are slightly pointed or acuminate .............ec-c0e0-. 22
21(20). Basimere with 3 slightly curved, prominent submarginal setae;
flagellomere VII with an irregularly angulate tuft of scales which
are kinked towards the apex.............6--. variatus (p. 72)
Basimere with 3 or 4 distinctly curved, prominent submarginal
setae; scales of the tuft on flagellomere VIII only gently curved...
macdonaldi (p. 61)
22(20). All scales in the tuft on flagellomere V acuminate; subapical lobe
of the basimere with the internal rod expanded apically and with
a Sealloped ARON: a viaicia a oe ee ee ee reidi (p. 66)
At most, some of these scales with a small short point; subapical
lobe of the basimere with the internal rod slightly expanded
apically but without a scalloped margin. ..........6+4-e-+00. 23.
23(22). Basimere with from 7 to 9 very long, strong, characteristically
curved, submarginal setae; palpus with external basal process
lonwor than SORWNONE Toca a os ob ss tw ees quadripalpis (p. 65)
Basimere with 3 slightly curved, prominent submarginal setae;
palpus with external basal process very inconspicuous, much
ghorter than seementl 4. lee es aculeatus (p. 50)
KEY TO SPECIES OF THE SUBGENUS LOPHOCERAOMYIA IN THAILAND -
FOURTH STAGE LARVAE!
a Siphon shorter than saddle; antennal shaft cylindrical, not con-
stricted beyond the insertion of antennal hair 1-A; mouthparts
adapted for DROCAUON, <p i. 2s 4 Mie aca eae ae aot curtipalpis (p. 76)
Siphon longer than saddle; antennal shaft constricted beyond
insertion of antennal hair 1-A; mouthparts not adapted for
DVOUAIION fic iice eles eine le 4 Ow ee ee se 0S 2
2(1). Head hair 4-C distinctly longer than the distance between the bases
of the pair; abdominal hair 2-VII single; thoracic hair 14-P
bifid or trifid (except in mammilifer and wilfredi). .........66.
(mammilifer subgroup) 3
! The larvae of lucaris and incomptus are unknown.
48
3(2).
4(3).
5(4).
6(5).
7(5).
8(7).
9(8).
10(8).
11(10).
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Head hair 4-C shorter than the distance between the bases of the
pair; abdominal hair 2-VII bifid (except in infantulus); thoracic
hair 14-P single (except in quadripalpis and infantulus)........
(fraudatrix group) 13
Comb consisting of 2 distinct types of scales, the posterior scales
tapering to a point, the anterior scales fan-shaped.......... y
minor (p. 96)
All comb scales fan-shaped, fringed with fine, subequal spines. ... A
Head hair 6-C always single, somewhat flattened. . . eukvines (p. 85)
Head hair 6-C branched, usually cylindrical...............e.6 3)
Siphon normally with 3 pairs of subventral tufts............... 6
Siphon normally with 4 pairs of subventral tufts ............... 7
Thoracic hair 14-P single; head hairs 16,17-C branched; abdominal
hair 6-III usually with 4 branches......... mammilifer (p. 93)
Thoracic hair 14-P bifid; head hairs 16,17-C single, relatively
long; abdominal hair 6-III usually with 3 branches............
bengalensis (p. 82)
Thoracic hair 8-P usually branched, always short and simple; ab-
OUT RI OSS DT nas oes case ese med caue eck pholeter (p. 1038)
Thoracic hair 8-P usually single, always long; abdominal hair
We ee ee ek ke be ee nck pk eek ss ee 8
Subventral tufts of the siphon fine, the length of the proximal tuft
less than the basal width of the siphon. ................0.-. 9
Subventral tufts of the siphon strong, the length of the proximal
tuft at least equal to the basal width of the siphon, usually
RO i ee fe es 10
Head hairs 16,17-C absent; thoracic hair 14-P usually bifid; integu-
ment of the siphon dark brown......... fuscosiphonis (p. 87)
Head hairs 16,17-C represented by minute, bifid spicules; thoracic
hair 14-P single; integument of siphon not exceptionally dark
Oe ee eG wilfredi (p. 110)
Thoracic integument covered with fine spicules; abdominal integu-
ment mostly smooth; median caudal filament not developed..... 11
Thoracic and abdominal integuments both covered with numerous
Spicules; median caudal filament short, but distinct......... 12
Head hair 1-C usually expanded on the basal half, then narrowing or
sometimes branched on the distal half; each pecten tooth fringed
with subequal lateral denticles............ ganapathi (p. 90)
Head hair 1-C never expanded on the basal half as indicated above;
each pecten tooth with from 1 to 3 proximal lateral denticles
much larger than the rést: 2... ek. peytoni (p. 100)
12(10).
13(2).
14(13).
15(14).
16(15).
17(16).
18(16).
19(18).
20(18).
Bram: Genus Culex in Thailand 49
Subventral tufts of the siphon finely pectinate; head hair 5-C bifid
or trifid; saddle with fine spicules on the distal margin........
spiculosus (p. 105)
Subventral tufts of the siphon simple; head hair 5-C usually with 3
or 4 branches; saddle with stronger spines on the distal margin...
traubi (p. 108)
Comb with a strongly differentiated posterior row of long scales;
head hair 5-C weak, its branches noticeably finer than those of
6-C and extending only a little past the anterior margin of the
heads 2 6) Bee i ee. alphus (p. 53)
Posterior row of comb scales not strongly differentiated; head
hair 5-C with branches about as stout as those of 6-C and
extending well past the anterior margin of the head.......... 14
Head hair 9-C much longer than 8-C, with spreading branches which
reach almost to the level of hair 7-C........ macdonaldi (p. 61)
Head hair 9-C normal, shorter, or not much longer than 8-C..... 15
Head hair 5-C with 3 or 4 branches........... cinctellus (p. peat
Head haiy S30 ified 74 6 Oe Bo ONG REY oe
Ab@ominal hair Tel Single <5. Or Pa ee. a ee 17
Abdominal hair’ 7-1 branched) oF es res, OE Ie ae 18
Thoracic hair 3-P with from 4 to 10 branches; 4-P bifid; thoracic
integument strongly spiculate........... rubithoracis (p. 69)
Thoracic hair 3-P usually single (occasionally bifid); 4-P single;
thoracic intesument Glabrous.020 0... eee ee es reidi (p. 66)
Thoracte hair-4>P usually single: 3543.8 Soe ON ais BS ws ws 19
Saoracic hair 4-P branched v5.0 GF ke ees ee Oe ee a ee 20
Pecten teeth, particularly the proximal ones, with strong basal
denticles; thoracic hair 3-P often single, pectinate............
nuadeedinis (p. 65)
Pecten teeth with a coarse fringe, but without strongly differentiated
basal denticles; thoracic hair 3-P usually bifid, simple.........
aculeatus (p. 50)
Thoracic hair 14-P single; 3-P single or at most bifid; head hair
4-C often with 3 or more branches; abdominal hair 2-VHI
Ta PO ee ee a, Ge oe a variatus (p. 72)
Thoracic hair 14-P bifid; 3-P often with 2 or 3 branches; head
hair 4-C often single, at most bifid; abdominal hair 2-VIII
GINO. ire Oe ek ee ek infantulus (p. 58)
FRAUDATRIX GROUP
FEMALE. In general, the scales on wing veins Rg and Rg are rather
50 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
broad and clavate, particularly towards the apex; otherwise inseparable from
members of the mammilifer group.
MALE. Head. Torus of the antenna normal, symmetrical, without a
protuberance on the inner surface; length of the palpus usually greater than
that of the proboscis, the palpus with distinct but small basal processes;
proboscis usually with long dorsal setae and a series of strong basoventral
bristles. Terminalia. Subapical lobe of the basimere usually with the in-
ternal and external leaflets well developed, subequal in size and shape;
lateral plate of the phallosome consisting of the dorsal process only.
LARVA. Head. Length of 4-C less than the distance between the
bases of the pair. Thorax. Hair 14-P single (except in quadripalpis and
infantulus). Abdomen. Hair 2-VIII bifid (except in infantulus); siphon with 4
pairs of subventral tufts (occasionally 9 individual tufts may be present);
central filament of the siphon valves well developed.
DISTRIBUTION. Species of the fraudatrix group are distributed
throughout the range of the subgenus. In Thailand, members of this group
are undoubtedly present in every changwat.
TAXONOMIC DISCUSSION. The characters outlined above for the
male and fourth stage larva clearly separate the fraudatrix group from the
mammilifer group. The one distinctive feature of the adult female, although
usually valid, is nebulous and should not be considered definitive.
Within the group, no definite species complexes such as those found
in the mammilifer group, are readily apparent; however, variatus does ex-
hibit extremely close relationships with macdonaldi, and with whartoni and
cubitatus from Malaya. Similarities also seem to be shared by quadripalpis
and aculeatus. Except for cinctellus and infantulus, both of which exhibit ©
narrow, pale, basal abdominal bands, the adult females are extremely similar
and cannot be conclusively recognized at the present time.
CULEX (LOPHOCERAOMYIA) ACULEATUS COLLESS 1965
(Figure 13)
Culex (Lophoceraomyia) aculeatus Colless 1965, J. med. Ent. 2: 286 (c*, 9,
L*),
The adult male may be recognized by the presence of 7 or less dorsal
scales on flagellomere V, by the presence of 3 curved submarginal setae on
the basimere of the male terminalia, and by the short external basal processes
of the male palpus. The fourth stage larva may be distinguished by abdominal
hair 7-I being branched, thoracic hair 4-P single, and by the absence of
exceptionally strong basal denticles on the pecten tooth.
FEMALE. Head. Proboscis with a pair of prominent basoventral
bristles and a pair of smaller setae between them. Thorax. Integument of
Figure 13. C. (Lophoceraomyia) aculeatus. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through X. “ii
Bram: Genus Culex
Cea Ly Z
II
C. (Lophoceraomyia) acu/eatus
in Thailand
o1
52 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
the pleuron uniformly pale, occasionally with dark, indistinct patterns; 1
lower mesSepimeral bristle present. Abdomen. Terga covered with dark
brown scales; sterna dull brown, but lighter than the terga.
MALE. Head. Proboscis with inconspicuous, short, dorsal setae
and approximately 10 fine basoventral setae; basal processes of the palpus
weak, fine and inconspicuous, shorter than segment I. Antenna. (Figure
13D). Flagellomere V with a tuft of 2 or 3 broad scales which are blunt or
slightly pointed apically followed by 2 or 3 broad, pointed scales of decreas-
ing length and 1 or 2 narrow, long, rather slender scales; VI with an internal
tuft of 10 or more short, dark, stout and twisted spines; VII with a similar,
but shorter tuft of spines and a tuft of 3 very broad spines; VIII with a tuft of
5 or more long, prominent spines which are gently curved distally; IX witha
series of straight, lanceolate, though narrow, leaflets; X with a tuft of from
3 to 5 strong, straight setae. Terminalia. (Figure 13C). Basimere with 3
gently curved submarginal setae inserted in a line; subapical lobe of the
basimere with the basal rods subequal in length with the internal rod moderate-
ly expanded; internal and external leaflets subequal in size, approximately
symmetrical, the internal leaflet with a fine apical filament; with up to 6
closely placed accessory setae which are subequal in length, difficult to
distinguish (only 3 clearly shown in illustration); distimere normal in shape,
with minute annulations on the convex surface of the apical fifth; dorsal
process of the lateral plate of the phallosome strongly recurved and pointed.
LARVA. (Figure 13A,B). Head. Antenna with a narrow dark basal
ring and progressively darker beyond insertion of hair 1-A; head hair 4-C
Single or with up to 4 branches, simple; 5,6-C bifid, pectinate, subequal in
length; 16,17-C represented by fine, relatively well developed spicules.
Thorax. Integument glabrous (occasionally appearing granulose or minutely
spiculose); 3-P usually bifid (occasionally single or trifid), shorter and
slenderer than 1,2-P, superficially simple, but with extremely fine lateral
barbs (not illustrated); 4,5,6-P single, pectinate; 7-P bifid or trifid, pecti-
nate; 8-P bifid, pectinate; 14-P single, simple. Abdomen. Integument
glabrous; hair 7-I branched; comb consisting of from 30 to 45 fan-shaped
scales arranged in a broad triangular patch; siphon index variable, ranging
from 7:1 to 10:1 (average, 7.8:1), the siphon occasionally with a broad
median dark band (not illustrated); 4 pairs of subventral tufts inserted in a
line on the siphon; individual tufts with 2 or 3 branches, their length ap-
proximately equal to the width of the siphon at the point of insertion; pecten
consisting of from 10 to 12 teeth restricted to the basal third or less of the
siphon; individual pecten tooth with a fine apical spine and from 7 to 10
lateral barbs which are progressively, but evenly, enlarged from apex to
base.
TYPE DATA. Holotype male of aculeatus (associated larval and pupal
skins and terminalia and antenna slide-mounted) from Perak, Malaya in the
Australian National Insect Collection, Canberra, Australia.
DISTRIBUTION. In THAILAND**, aculeatus has been studied from:
Chiang Mai, Krung Thep, Lampang, Nakhon Si Thammarat, Narathiwat,
Ranong, Songkhla, Tak, Trang, and Yala. In MALAYA the species was
recorded by Colless (1965) from Bukit Tunggal, Perak (the type locality)
and Ampang and Ulu Gombak, Selangor.
During this study the following material was examined: 1 adult female
(a paratype); 1 adult male with associated larval and pupal skins (a paratype),
Bram: Genus Culex in Thailand 53
as well as 8 males with their associated terminalia and antennae, and 23
fourth stage larvae.
TAXONOMIC DISCUSSION. As indicated previously, aculeatus and
quadripalpis demonstrate close affinity in all stages but may be distinguished
by anatomical features of the adult male and the fourth stage larva.
BIOLOGY. In Thailand, larvae have been collected from slow flowing
streams, rock pools, ground pools, stream pools, and Seepage areas. Biol-
ogy of the adults is virtually unknown, but they have been taken by sweeping
foliage in the jungle and at light traps. Colless (1965) has suggested that it
is possible that the females may occasionally attack man in jungle foliage.
CULEX (LOPHOCERAOMYIA) ALPHUS COLLESS 1965
(Figure 14)
Culex (Lophoceraomyia) alphus Colless 1965, J. med. Ent. 2: 283 (o*, 9, L*).
The adult female may be recognized by the presence of a ventral
transverse row of 5 or more strong bristles at the base of the proboscis
(some of which project laterally). The adult male exhibits a distinctive scale
tuft on flagellomere V and abdominal sternum VII is covered with white scales
which contrast with the dark integument and scaling of the other sterna. In
the fourth stage larva the comb possesses a strongly differentiated posterior
row of long scales and head hair 5-C is weak, its branches noticeably finer
than those of 6-C and extending only a little past the anterior margin of the
head.
FEMALE. Head. Proboscis with 3 pairs of prominent basoventral
bristles and a pair of smaller setae between them. Thorax. Integument of
pleuron uniformly dark brown; 1 lower mesepimeral bristle present; anterior
surface of hind femur completely pale proximally, dark distally. Abdomen.
Terga dark, with a slight mixture of light and dark scales on the lateral
margins of terga VI and VU; sterna completely white.
MALE. Head. Proboscis with numerous hairs on the distal 2/3 of
the dorsal surface and with about 10 basoventral bristles present. Antenna.
(Figure 14D). Flagellomere V with a tuft of broad, bluntly rounded scales
which extend to IX followed by 4 or 5 narrow, short, pointed scales, and then
2 to 4 longer scales which have a rather narrow base but are noticeably ex-
panded at the apex; VI with an internal tuft of 10 or more short, dark, stout
and twisted spines; VII with a similar, but shorter tuft of spines; VID witha
tuft of 6 or more long, prominent spines which are gently curved distally; IX
with a tuft of sharply pointed, lanceolate leaflets; X with from 3 to 5 strong,
straight setae. Abdomen. Sternum VII with white scaling which sharply
contrasts with the general dark color of the other abdominal segments.
Terminalia. (Figure 14C, after Colless 1965). Basimere with 4 long and
apically curved submarginal setae; subapical lobe of the basimere with the
rods subequal in length, the internal rod slightly expanded apically; internal
and external leaflets broad, the internal leaflet asymmetrical; 6 accessory
setae blade-like and of varying lengths, 1 of which exhibits a serrate edge;
distimere normal, without annulations; dorsal process of the lateral plate of
the phallosome short, strongly recurved, and directed inwards as well as
dorsally.
54 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.14
we)
‘
SAH:
. \ MM Wf
\\\ \ WA
\i WY
SS
Ss
>
C. (Lophoceraomyia ) alphus
Bram: Genus Culex in Thailand 55
LARVA. (Figure 14A,B). Head. Antenna slightly darker than the
head capsule and with a narrow, dark basal ring and dark beyond insertion of
hair 1-A; head hair 4-C with from 3 to 6 branches, simple, its length con-
siderably less than the distance between the bases of the pair; 5-C bifid,
pectinate, its branches noticeably finer than 6-C, extending only a little past
the anterior margin of the head; 6-C bifid or trifid, pectinate, considerably
longer and broader than 5-C, more than half its length projecting beyond the
anterior margin of the head. Thovax. Integument glabrous; hair 3-P con-
siderably shorter than 1,2-P, with 6 to 8 branches; 4-P bifid, pectinate,
inserted close to 3-P, weak and smaller than the other prothoracic hairs;
5,6-P single, pectinate; 7-P trifid, pectinate; 8-P bifid, pectinate; 14-P
single or bifid, simple; 0-P dendritic, very large, usually longer than 4-C.
Abdomen. Integument glabrous; comb consisting of approximately 20 fan-
shaped scales arranged in 2 irregular rows; the scales of the posterior row
elongated, with long basal attachments, those of the anterior row short;
siphon index approximately 7:1; 4 pairs of subventral tufts inserted in a line
on the siphon; individual tufts with from 2 to 6 branches, their length greater
than the width of the siphon at the point of insertion; pecten consisting of
from 8 to 10 teeth restricted to the basal third of the siphon; individual pecten
tooth consisting of a strong spine without lateral barbs, but with a very fine
lateral fringe.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia and antenna slide-mounted) from Kg. Sijangkang, Selangor, Malaya
in the Australian National Insect Collection, Canberra.
DISTRIBUTION. In THAILAND**, a single male has been collected
from Narathiwat at a light trap. Colless (1965) reported examining specimens
from SINGAPORE and Selangor, MALAYA and NORTH BORNEO.
In addition to the specimen from Thailand, the author has examined
the paratype series in the British Museum, and it is from these specimens
that the above description has been prepared.
TAXONOMIC DISCUSSION. This species is extremely characteristic
and there should be no difficulty in identifying either the adult male (with or
without antenna mounted) or the fourth stage larva.
BIOLOGY. Larvae have been collected on one occasion from a shaded
well. Males have been collected at light traps and resting in secondary jungle.
CULEX (LOPHOCERAOMYIA) CINCTELLUS EDWARDS 1922
(Figure 15)
Lophoceratomyia taeniata Leicester 1908, Cul. Malaya: 127 (preoccupied by
Wiedemann 1828).
Culex (Lophoceratomyia) cinctellus Edwards 1922, Indian J. med. Res. 10:
Figure 14. C. (Lophoceraomyia) alphus. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral
aspect of the terminal abdominal segments of the fourth stage
hoo larva; C, dorsal aspect of the basimere, lateral aspect of the
phallosome of the male terminalia (after Colless 1965); D,
lateral aspect of male antennal flagellomeres V through X.
56 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
281 (new name for /aeniata Leicester); Barraud 1924, Indian J. med.
Res. 12: 42 («*); Barraud 1934, Fauna Brit. India, Diptera 5: 366
(adult only).
Culex (Lophoceraomyia) cinctellus Edwards: Colless 1965, J. med. Ent. 2:
277 (o*, 2, L*); Delfinado 1966, Mem. Amer. ent. Inst. 7: 103 (o*).
The adult female may be recognized by the basal banding of the ab-
dominal terga and the predominantly dark decumbent scales of the vertex.
The male is distinctive in that the submarginal setae of the basimere are in-
serted in a group rather than in a straight line. The fourth stage larva is
separated from other members of the fraudatrix group by the presence of a
3 or 4 branched head hair 5-C.
FEMALE. Head. Decumbent scales of the vertex dark brown, be-
coming pale immediately adjacent to the orbital line. Thovax. Scutum and
midlobe of the scutellum sparsely covered with reddish brown scales; 1
strong lower mesepimeral bristle present; anterior surface of the hind femur
pale, with a very broad, dark apical band which extends proximally as a
narrow dark stripe along the dorsal border. Abdomen. Terga dark brown,
with narrow, complete pale basal bands; sterna pale brown.
MALE. Head. Proboscis with rather long dorsal setae and a linear
series of strong basoventral setae; palpus with very short, inconspicuous
basal processes. Antenna. (Figure 15D). Flagellomere V with a promi-
nent tuft of from 3 to 5 dark, blade-like, sharply pointed scales which ex-
tend to VIII, accompanied by 1 to several strong, very long setae; VI with an
internal tuft of 10 or more short, dark, stout and twisted spines; VII with a
similar, but shorter, tuft of spines and a supplementary tuft of 3 longer,
broader spines; VIII with a tuft of 7 or more long, prominent spines which
are gently curved; IX with a tuft of sharply pointed, lanceolate leaflets as
well as several strong, straight setae; X with a tuft of from 2 to 5 strong,
straight setae. Terminalia. (Figure 15C). Basimere with 14 or more strong
submar ginal setae inserted in a conspicuous patch; subapical lobe of the
basimere with the rods subequal in length and thickness, the internal rod ex-
panded distally; internal and external leaflets broad, the apex of the internal
leaflet with an extended point; accessory setae rather broad and prominent;
distimere noticeably expanded throughout, with minute annulations at the
apex; dorsal process of the lateral plate of the phallosome angulate and sharply
pointed.
LARVA. (Figure 15A,B). Head, Antenna with a narrow, dark basal
band and more darkly pigmented beyond insertion of hair 1-A; head hair 4-C
simple, with from 2 to 4 branches; 5-C 3 or 4 branched, pectinate; 6-C bifid,
pectinate; 16,17-C represented by minute spicules. Thorax. Integument
superficially glabrous, but with some very fine spiculation on the anterio-
lateral surface of the prothorax (not illustrated); hair 3-P considerably shorter
Figure 15. C. (Lophoceraomyia) cinctellus. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male antennal
flagellomeres V through X.
: =~
Bram: Genus Culex in Thailand
SSS
SS
SSS
NSS
y
WW
WY ZS
SSS
Wy
C. (Lophoceraomyia) cinctellus
HOY MNT
58 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
and slenderer than 1,2-P, with from 2 to 4 pectinate branches; 4-P bifid,
pectinate; 5,6-P single, pectinate; 7-P with 3 or 4 branches, pectinate; 8-P
bifid, pectinate; 14-P single, simple. Abdomen. Integument glabrous; comb
consisting of from 40 to 50 fan-shaped scales arranged in a broad, triangular
patch; siphon index approximately 8:1; 4 pairs of subventral tufts inserted in
a line on the siphon; individual tufts 2 or 3 branched, simple, their length
less than the width of the siphon at the point of insertion; pecten consisting of
from 8 to 11 teeth restricted to the basal fourth or less of the siphon; individual
pecten tooth with a fine distal spine and from 11 to 14 very fine lateral barbs.
TYPE DATA. Lectotype male (terminalia mounted on a celluloid
point) of taeniata from Kuala Lumpur, Malaya in the British Museum.
DISTRIBUTION. In THAILAND**, this species has been collected
from Chon Buri and Khon Kaen. The species is also known from MALAYA,
INDIA, BORNEO, INDONESIA, HAINAN, and the PHILIPPINES.
Twelve males and their associated terminalia and antennae have been
studied from Thailand. One male and 1 female with their associated larval
skins have been seen from Malaya and it is from these skins that the larval
description was prepared.
TAXONOMIC DISCUSSION. Due to the pale, basal abdominal banding,
cinctellus appears to have its closest affinity to infantulus. The male termi-
nalia and antennal modifications of the 2 species are quite distinctive; how-
ever, Delfinado (1966) has pointed out that cinctellus and fulleri (Ludlow)
from the Philippines cannot be separated on the basis of the male antenna and
terminalia, but fullevi does not exhibit the pale basal bands on the abdominal
terga.
BIOLOGY. In Thailand, adult males have been collected only at light
traps and resting in a bamboo forest during the months of October and
November. Colless (1959a) has suggested that adults of this species feed
largely on birds, but occasionally man serves as a host. In Malaya larvae
were collected in shaded ground pool waters and stream margins in or near
forests.
CULEX (LOPHOCERAOMYIA) INFANTULUS EDWARDS 1922
(Figure 16)
Culex (Lophoceratomyia) infantulus Edwards 1922, Indian J. med. Res. 10:
287 (co); Baisas 1935, Philipp. J. Sci. 57: 174 («*); Edwards 1935,
Bull. ent. Res. 26: 131 (L); Causey 1937, Amer. J. Hyg. 25: 416
(distribution); LaCasse and Yamaguti 1950, Mosq. Fauna Japan and
Korea: 197 (o*, 9*, L*, P*),
Culex (Lophoceratomyia) parainfantulus Menon 1944, J. Malar. Inst. India 5:
389 (o*, 9*); Mattingly 1949, Proc. R. ent. Soc. Lond. (B) 18: 224
(synonymy).
Culex (Lophoceraomyia) infantulus Edwards: Bohart 1945, Navmed 580: 75
(o, L); Bohart and Ingram 1946, Navmed 1055: 73 (o'*, 9, L*, P*);
Iyengar and Menon 1955, Bull. ent. Res. 46: 10 (L*); Peters and
Dewar 1956, Indian J. Malar. 10: 46 (L); Delfinado 1966, Mem. Amer.
ent. Inst. 7: 106 (c*).
The presence of pale basal abdominal bands clearly separates the adult
Bram: Genus Culex in Thailand 59
female from most other members of the subgenus. The male is recognized
by the reduced number of modified setae on the antennal flagellum and the
reticulate dorsal process of the phallosome. The fourth stage larva exhibits
thoracic hair 3-P with 2 or 3 branches, 4-P branched, and 14-P bifid.
FEMALE. Thorax. Integument of the pleuron dark brown; 1 strong
lower meSepimeral bristle present; wings with the outstanding scales of
veins R2 and R3 clavate, but not as broad as in most other species of the
fraudatrix group. Abdomen. Terga and sterna dark brown, with narrow,
complete pale basal bands (in some specimens these bands may be represent-
ed by a single row of pale scales).
MALE. Head. Proboscis without exceptionally long dorsal or ventral
hairs; palpus with very short, inconspicuous basal processes. Antenna.
(Figure 16D). Flagellomeres V, VI, IX, and X without modified setae or
scales; VII with a tuft of 1 or 2 very broad, dark setae and often with several
narrow, pointed setae of similar length; VIII with from 3 to 5 broad, long and
gently curved setae. Terminalia. (Figure 16C). Basimere with 3 fine sub-
marginal setae inserted in a straight row; subapical lobe of the basimere
with the internal, central, and external rods subequal in length and width,
hooked apically; accessory setae reduced in size and number; internal leaflet
very short, bladelike, external leaflet somewhat broader; distimere normal;
dorsal process of the lateral plate of the phallosome rather broad, gently
curved and pointed, exhibiting a distinctive reticular pattern over its entire
surface.
LARVA. (Figure 16A,B). Head. Antenna with a narrow, dark basal
band and somewhat darker beyond insertion of hair 1-A; head hair 4-C single,
simple, its length approximately equal to or slightly shorter than the distance
between the bases of the pair; 5,6-C bifid, pectinate, subequal in length;
16,17-C represented by minute, short spicules which are easily overlooked
even in good preparations. Thovax. Integument glabrous; hair 3-P bifid,
pectinate, the branches slenderer and shorter than 1,2-P; 4-P bifid, pecti-
nate; 5,6-P single, pectinate; 7-P trifid, pectinate; 8-P bifid, pectinate; 14-P
bifid, simple. Abdomen. Integument glabrous; hair 7-I branched; 2-VIII
single; comb consisting of from 30 to 50 fan-shaped scales arranged ina
broad triangular patch; siphon index variable, ranging from 9:1 to 11. 6:1
(average, 10:1); arather broad median dark band occasionally present on the
siphon (not illustrated), if present not as dark as the basal ring; 4 pairs of
subventral tufts inserted in a line on the siphon; individual tufts usually bifid
(occasionally trifid), their length equal to or less than the width of the siphon
at the point of insertion; pecten consisting of from 12 to 17 teeth restricted to
the basal fourth or less of the siphon; individual pecten tooth with a fine distal
Spine and from 5 to 8 lateral barbs.
TYPE DATA. Holotype male (terminalia mounted and attached to pin)
of infantulus from Hong Kong in the British Museum. The type locality of
parainfantulus is Trivandrum, Travancore, India, but the location of the type
specimen is unknown.
DISTRIBUTION. In THAILAND, infantulus has been collected from:
Ayutthaya, Chanthaburi, Chiang Mai, Chon Buri, Nakhon Ratchasima, Nan,
Pathum Thani, Phattalung, Prachin Buri, Ranong, Satun, Songkhla, Tak,
Trang, and Udon Thani. The species has also been recorded from CHINA,
JAPAN, RYUKYU-RETTO, INDIA, CEYLON, NEPAL, MALDIVE ISLANDS,
INDOCHINA, JAVA, and the PHILIPPINES. Specimens have also been ex-
60
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
wm
}
H)
WW,
OUR
Sy
CH
SEEING
C. (Lophoceraomyia/) infantulus
Fig.16 -
14
id
Bram: Genus Culex in Thailand 61
amined in the British Museum from Rangoon, BURMA** (xii 1962, P. F.
Mattingly).
During this study 9 individual rearings have been examined as well as
11 males, 2 females, and 40 larvae.
TAXONOMIC DISCUSSION. The pale basal abdominal bands of the
adult female distinguish this species from all other members of the fraudatrix
group except cinctellus; these species may be distinguished, however, on the
basis of the narrow decumbent scales of the vertex which are uniformly pale
in infantulus and predominantly dark in cinctellus. The male of infantulus is
separated from all other members of the subgenus by the absence of special-
ized scales or setae on antennal flagellomeres V, VI, IX, and X, and by the
reticular pattern on the dorsal process of the phallosome. The larva dem-
onstrates its closest affinity to variatus but may be separated on the basis of
head hair 4-C which is usually single in znfantulus and branched in variatus,
and thoracic hairs 3,14-P which are usually bifid in infantulus and single in
vaviatus; the larva of infantulus is the only member of the fraudatrix group in
which abdominal hair 2-VIII is single.
BIOLOGY. In Thailand, larvae of this species have been collected
from a wide variety of both ground water and container habitats. Collections
have been made from ground pools, ditches, pools in dry stream beds, tree
holes, a jar, rock pools, bamboo internodes, and small crab holes next toa
stream. Nothing is known of the adult biology, but the females have never
been reported biting man.
CULEX (LOPHOCERAOMYIA) MACDONALDI COLLESS 1965
(Figure 17)
Culex (Lophoceraomyia) macdonaldi Colless 1965, J. med. Ent. 2: 276 (c*,
?, L*); Delfinado 1966, Mem. Amer. ent. Inst. 7: 108 («*, L*, P*).
The adult male is recognized by the long basal processes of the palpus
and the nature of antennal flagellomere V. The larva is immediately recog-
nized on the basis of head hair 9-C which is much larger than 8-C.
FEMALE. Specimens not available for study.
MALE. Head. Proboscis with rather long, straight dorsal setae and
several fine basoventral setae; palpus with long basal processes which extend
beyond the apex of palpal segment II. Antenna. (Figure 17D). Flagellomere
V with from 9 to 12 long, dark, blunt, striated scales dorsally and approxi-
mately 8 shorter, broad, striated, pointed scales which appear darker on
their basal halves; VI with a tuft of 13 or more short, stout, dark and twisted
spines; VII with a similar but shorter tuft; VIII with a tuft of 5 or more broad,
long spines which are gently curved; IX with from 3 to 5 finely pointed lan-
ceolate leaflets as well as several stout setae; X with an internal tuft of 3 or 4
Figure 16. C. (Lophoceraomyia) infantulus. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the male terminalia; D, lateral aspect of
male antennal flagellomeres V through X.
athe
62
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
4 Andi)
Wy PT
y y
Wy
Wy
\ MN | ie
C. (Lophoceraomyia ) macdonaldi
rig. hz:
Bram: Genus Culex in Thailand 63
straight, strong, long setae. Thovax. Integument of the pleuron pale brown;
1 strong lower mesepimeral bristle present; anterior surface of the hind
femur predominantly dark, with a small basoventral pale area. Abdomen.
Terga dark, sterna paler, Terminalia. (Figure 17C). Basimere with 4 or 5
strong, submarginal setae inserted in a straight row; subapical lobe of the
basimere with the basal rods subequal in length and thickness, the central rod
somewhat expanded distally; accessory setae narrow, indistinct; the internal
and external leaflets subequal in size and shape; distimere expanded and
annulate on the apical half; dorsal process of the lateral plate of the phallo-
some rather short, pointed, and sharply bent, the posterior angle smooth.
LARVA. (Figure 17A,B). Head. Antenna with a narrow, dark, basal
band and darkly pigmented beyond insertion of hair 1-A; head hair 4-C short,
with from 2 to 5 branches; 5, 6-C bifid, pectinate, subequal in length; 9-C
with from 4 to 6 long branches, much longer than 8-C and almost reaching to
the base of 7-C; 16,17-C represented by minute spicules. Thovax. Integu-
ment glabrous; 3-P considerably shorter and slenderer than 1,2-P, with from
2 to 4 branches; 4-P bifid, pectinate; 5,6-P single, pectinate, 5-P branched
at the apex; 7-P trifid, pectinate; 8-P bifid, pectinate; 12-P with 3 short
branches (in contrast to the usual single, very long branch); 14-P single,
simple. Abdomen. Integument glabrous; comb consisting of from 35 to 45
fan-shaped scales arranged in a broad triangular patch; siphon index variable,
ranging from 7:1 to 9:1; 4 pairs of subventral tufts inserted in a line on the
siphon; individual tufts either single or with up to 4 branches, simple, their
length less than the basal width of the siphon; pecten consisting of from 8 to
10 teeth restricted to approximately the basal 1/4 of the siphon; individual
pecten tooth with a fine distal spine and from 7 to 10 fine lateral barbs, the
proximal 1 or 2 barbs considerably more prominent than the other lateral
barbs.
TYPE DATA. Holotype male (antennae, terminalia, and associated
larval and pupal skins slide-mounted) from Singapore in the Australian Na-
tional Insect Collection, Canberra.
DISTRIBUTION. In THAILAND**, macdonaldi has been collected
from: Chanthaburi, Chiang Mai, Chon Buri, Krung Thep, Narathiwat, Sakon
Nakhon, and Thon Buri. The species has also been reported from SINGA-
PORE, MALAYA, ASSAM and the PHILIPPINES.
Eleven males with their associated terminalia and antennae have been
examined as well as 2 larvae and a paratype male with its associated larval
and pupal skins from Singapore.
TAXONOMIC DISCUSSION. Although this species ee its
closest affinity to variatus, it is easily distinguished on the basis of charac-
ters presented in the first paragraph.
BIOLOGY. Larvae have been collected from fresh water just above
tidal limits and in Thailand from rock pools. Adults have been collected at
light traps, but virtually nothing is known of the biology of the species.
Figure 17. C. (Lophoceraomyia) macdonaldi. A, dorsoventral aspect of
the head and prothorax of the fourth stage larva; B, lateral
—— aspect of the terminal abdominal segments of the fourth stage
larva; C, dorsal aspect of the basimere, lateral aspect of the
phallosome of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through x.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
64
omyia) quadripalpis
C.(Lophocera
Bram: Genus Culex in Thailand | 65
CULEX (LOPHOCERAOMYIA) QUADRIPALPIS (EDWARDS) 1914
(Figure 18)
Lophoceratomyia sylvestris Leicester 1908, Cul. Malaya: 125 (not Culex
sylvestris of Theobald 1901); Edwards 1922, Indian J. med. Res. 10:
473 (synonymy).
Lophoceratomyia quadripalpis Edwards 1914, Bull. ent. Res. 5: 80 (¢).
Culex (Lophoceratomyia) quadripalpis (Edwards): Edwards 1922, Indian J.
med. Res. 10: 473 (taxonomy).
Lophoceratomyia roubaudi Borel 1926, Arch. Insts. Pasteur Indo-Chine 3-4:
38 (o'*, 2, L*); Borel 1927, Bull. Soc. Pat. exot. 20: 25 (c*, ?, L*);
Borel 1930, Monogr. Coll. Soc. Pat. exot. 3: 362 (o*, 9, L*); Ed-
wards 1932, in Wytsman, Genera Insect. fasc. 194: 197 (synonymy).
Culex (Lophoceraomyia) quadripalpis (Edwards): Mattingly 1949, Proc. R.
ent. Soc. Lond. (B) 18: 227 (L*, taxonomy); Colless 1965, J. med.
Ent; .2: 265 (o*, 9, 1%),
The adult male may be recognized by the presence of from 1 to 4
broad, dorsal scales on flagellomere V, by the 7 or 8 very long, curved sub-
marginal setae on the basimere of the terminalia, and by the relatively long
external basal processes of the palpus. The fourth stage larva is rather in-
distinctive, but abdominal hair 7-I is branched, thoracic hair 4-P is single,
and the proximal pecten teeth have strong basal denticles.
FEMALE. Thorax. Integument of the pleuron grayish to pale brown;
1 lower mesepimeral bristle present. Abdomen. Terga and sterna dark
brown.
MALE. Head. Proboscis with conspicuous dorsal setae and with 10
or more strong basoventral bristles; palpus with the basal processes extend-
ing approximately to the apex of palpal segment II. Antenna. (Figure 18D).
Flagellomere V with a tuft of from 2 to 4 broad scales which are usually
pointed (occasionally blunt or rarely notched) and extend almost to the tuft on
IX, these are followed by from 3 to 5 shorter, pointed scales; VI with an in-
ternal tuft of 10 or more short, dark, stout and twisted spines; VII witha |
similar, but shorter tuft of spines; VII with a tuft of 7 or more long, promi-
nent spines which are gently curved distally; [IX with a tuft of sharply pointed,
lanceolate leaflets as well as several strong, straight setae; X with a tuft of
from 3 to 5 strong, straight setae. Terminalia. (Figure 18C). Basimere
with from 7 to 9 strong, characteristically curved submarginal setae; sub-
apical lobe of the basimere with the basal rods subequal in length, the internal
rod slightly expanded; internal and external leaflets broad, the external leaf-
let smaller than the asymmetrical internal leaflet; 6 accessory setae closely
placed, equal in length and difficult to distinguish; distimere normal in shape,
with minute annulations on the convex surface of the apical fifth; dorsal
Figure 18. C. (Lophoceraomyia) quadripalpis. A, dorsoventral aspect of
the head and prothorax of the fourth stage larva; B, lateral
<< aspect of the terminal abdominal segments of the fourth stage
larva; C, dorsal aspect of the basimere, lateral aspect of the
phallosome of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through X.
66 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
process of the lateral plate of the phallosome strongly recurved and pointed,
the posterior angle of which usually exhibits from 1 to several very small
teeth.
LARVA. (Figure 18A,B). Head. Hair 4-C simple, usually single,
but sometimes exhibiting up to 3 branches; 5, 6-C bifid, pectinate, subequal in
length; 16, 17-C represented by rather well developed spines. Thorax. Integu-
ment glabrous; hair 3-P shorter and slenderer than 1,2-P, usually single,
pectinate (occasionally bifid); 4,5, 6-P single, pectinate (4-P occasionally.
bifid); 7-P trifid, pectinate; 8-P bifid, pectinate; 14-P bifid, simple, some-
times single on one side. Abdomen. Integument glabrous; hair 7-I branched;
comb consisting of from 30 to 40 fan-shaped scales arranged in a broad,
somewhat triangular patch; siphon index variable, ranging from 7. 4:1 to 10:1
(average, 9.1:1); a broad, dark median band usually present on the siphon; 4
pairs of subventral tufts inserted in a line on the siphon; individual tufts 2 to
5 branched, simple, their length equal to, or slightly greater than the width
of the siphon at the point of insertion; pecten consisting of from 10 to 14 teeth
restricted to the basal third or less of the siphon; individual pecten tooth with
a fine apical spine and from 5 to 8 lateral barbs, the proximal 1 or 2 barbs
considerably stronger and more prominent than the distal barbs.
TYPE DATA. Holotype male (terminalia and antennae slide-mounted)
of quadripalpis from Sarawak in the British Museum. Lectotype male (termi-
nalia slide-mounted) of sylvestris from Kuala Lumpur in the British Museum.
The type locality of rvoubaudi is Terres Rouges, Cochin-China, Indochina, but
the location of the type specimen is unknown.
DISTRIBUTION. In THAILAND**, guadripalpis has been collected
from: Krung Thep, Nakhon Nayok, Nakhon Ratchasima, Narathiwat, Satun,
Songkhla, and Tak. This species has also been recorded from SINGAPORE,
MALAYA, BORNEO, and INDOCHINA.
During this study the following material has been examined: 7 fe-
males with their associated larval and pupal skins; 11 males, 5 with their
associated larval and pupal skins; and 56 fourth stage larvae.
TAXONOMIC DISCUSSION. In both the adult male and the fourth stage
larva, quadripalpis demonstrates its closest affinity with aculeatus. In the
male, these species may be distinguished on the basis of the submarginal
setae of the baSimere which are 7 or 8 in number in quadripalpis and only 3
in aculeatus; in the fourth stage larva the shape of the pecten tooth is diag-
nostic, having at least one distinctive basal denticle in quadripalpis, and
thoracic hair 3-P usually single in quadripalpis but usually bifid in aculeatus.
BIOLOGY. In Thailand, larvae have been collected from a variety of
ground water habitats including stream pools, stream margins, pits, seepage
areas, and elephant hoof prints. Colless (1965) also collected larvae in
Malaya from fresh water ground pools and rock pools under forest shade and
usually associated with a considerable amount of decaying vegetable matter.
Little is known of the adult biology, but the females have been reported oc-
casionally attacking man in forest areas and adults have also been taken rest-
ing on forest vegetation.
CULEX (LOPHOCERAOMYIA) REIDI COLLESS 1965
(Figure 19)
Culex (Lophoceraomyia) reidi Colless 1965, J. med. Ent. 2: 279 (o*, 9, L*);
Delfinado 1966, Mem. Amer. ent. Inst. 7: 113 (c*).
Bram: Genus Culex in Thailand 67
Culex (Lophoceratomyia) quadripalpis of Edwards 1928 (not Edwards 1914),
Bull. ent. Res. 18: 276 (key); Edwards and Given 1928, Bull. ent.
Res. 18: 352 (L).
The adult male may be recognized by the presence of a tuft of 5 or 6
broad, but pointed scales on flagellomere V and by the internal rod on the
subapical lobe of the basimere being expanded distally with a scalloped mar-
gin. The fourth stage larva is identified by abdominal hair 7-I single,
thoracic hair 3-P usually single, 4-P single, and thoracic integument
glabrous.
FEMALE. Head. Decumbent scales of the vertex predominantly
broad, with a triangular median patch of narrow dark scales, the broad
scales being quite pale on the lateral margins; erect scales uniformly dark
brown. Thorax. Integument of the pleuron pale brown; 1 strong lower
mesepimeral bristle present; anterior surface of hind femur completely dark
on the distal half, predominantly pale on the basal half, but with a narrow
dark stripe along the dorsal border. Abdomen. Terga dark scaled, fre-
quently with paler, bronzy basolateral spots on segments VI and VII, but
occasionally extending to segment II; sterna dull brown.
MALE. Head. Proboscis with rather long dorsal setae and with
several long, basoventral bristles; palpus with long basal processes whose
apices usually extend beyond the apex of palpal segment Il. Antenna. (Fig-
ure 19D). Flagellomere V with a tuft of 5 or 6 broad, pointed scales which
extend to VIII, and occasionally several straight, long setae; VI with an in-
ternal tuft of 15 or more short, stout, dark and twisted spines; VII witha
similar, but smaller tuft of spines; VII with a tuft of slightly longer, smooth-
ly curved setae; [IX with a tuft of from 2 to 5 finely pointed, lanceolate leaf-
lets; X with a tuft of 4 strong, straight setae. Terminalia. (Figure 19C).
Basimere with from 3 to 5 submarginal setae inserted in a straight row; in-
dividual seta somewhat expanded and sharply angled at a little past the center;
subapical lobe of the basimere with the internal rod expanded apically and
with a scalloped margin; central and external rods subequal in length and
thickness, hooked; approximately 6 accessory setae present, apparently 2
setae and 4 blades; internal and external leaflets subequal in size, the in-
ternal leaflet asymmetrical; distimere expanded on the apical third and
minutely annulate on the convex margin; dorsal process of the lateral plate
of the phallosome short, sharply angled and pointed, the posterior angle of
which is smooth. .
LARVA. (Figure 19A,B). Head. Antenna with a narrow, dark basal
band and in some specimens, darker beyond insertion of hair 1-A; head hair
4-C simple, with from 1 to 4 branches, inserted relatively close to 6-C; 5, 6-C
bifid, pectinate, subequal in length; 16, 17-C represented by rather well de-
veloped spicules. Thorax. Integument glabrous; 3-P shorter and slenderer
than 1,2-P, usually single, pectinate (occasionally bifid); 4,5,6-P single,
pectinate; 7-P bifid or trifid, pectinate; 8-P bifid, pectinate; 14-P single,
Simple. Abdomen. Integument glabrous; hair 7-I single; comb consisting of
from 35 to 45 fan-shaped scales arranged in a broad, triangular patch; siphon
index variable, ranging from 5. 5:1 to 6.7:1 (average, 6.2:1), the siphon oc-
casionally with a broad, slightly darker median band (not illustrated); 4 pairs
of subventral tufts inserted in a line on the siphon; individual tufts with from
2 to 5 branches, simple, their length slightly greater than the width of the
siphon at the point of insertion; pecten consisting of from 11 to 13 teeth
restricted to approximately the basal third to fourth of the siphon; individual
68
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Wy
A) ire
it t
an
*
ANY
C. (Lophoceraomyia/ reidi
Bram: Genus Culex in Thailand 69
pecten tooth with a fine apical spine and from 8 to 12 lateral barbs, the
proximal barbs somewhat more prominent than the distal ones.
TYPE DATA. Holotype male (terminalia slide-mounted) and a series
of specimens reared from the same egg raft from Singapore in the Austral-
ian National Insect Collection, Canberra.
DISTRIBUTION. Specimens have been collected in THAILAND** from:
Krung Thep (Bangkok) and Thon Buri. The species is also known from
SINGAPORE and Selangor, MALAYA (Colless 1965), and from the PHILIP-
PINES (Delfinado 1966). |
During this study 2 males with their associated terminalia and
antennae and 5 larvae have been studied from Thailand, as well as a paratype
male and female with associated larval and pupal skins from Singapore.
TAXONOMIC DISCUSSION. The adult female demonstrates closest
affinity to variatus and macdonaldi, and no consistently distinctive characters
have been found which can be used to separate these species. The adult male
is separated from variatus and macdonaldi on the basis of the tuft on flagel-
lomere V and is separated from quadripalpis and aculeatus on the basis of the
internal rod on the subapical lobe of the basimere. The larva demonstrates
close similarity to rubithoracis but the thoracic chaetotaxy is distinctive.
BIOLOGY. Colless (1965) reported collecting larvae from the tidal
zone in Shaded pools, axils of Nipa palms, and in crab holes. In Thailand,
larvae have been collected from running water in a ditch. The adult females
have been reported attacking man, but the normal host complex remains un-
known.
CULEX (LOPHOCERAOMYIA) RUBITHORACIS (LEICESTER) 1908
(Figure 20)
Lophoceratomyia rubithoracis Leicester 1908, Cul. Malaya: 119 (<, 9);
Edwards 1913, Bull. ent. Res. 4: 236 (taxonomy).
Culex (Lophoceratomyia) rubithoracis (Leicester): Barraud 1934, Fauna
Brit. India, Diptera 5: 367 (co, ?); Hsiao and Bohart 1946, Navmed
1095: 25 (distribution); Bohart 1946, Navmed 961: 18 (key); LaCasse
and Yamaguti 1950, Mosq. Fauna Japan and Korea: 192 (o'*, 9*, L*¥, P*);
Hara 1957, Jap. J. exp. Med. 27: 57 (?*).
Culex (Lophoceraomyia) rubithoracis (Leicester): Dantis 1947, Mon. Bull.
Philipp. Hlth. Serv. 23: 255 (L, P); Mattingly 1949, Proc. R. ent.
Soc. Lond. (B) 18: 224 (L*); Colless 1959, Proc. R. ent. Soc. Lond.
(B) 28: 114 (L*); Lien 1962, Pacif. Ins. 4: 632 (distribution);
Colless 1965, J. med. Ent. 2: 281 (o*, 9, L*); Delfinado 1966, Mem.
Amer. ent. Inst. 7: 113 (<*).
The absence of a lower mesSepimeral bristle and the reddish brown tint
Figure 19. C. (Lophoceraomyia) reidi. A, dorsoventral aspect of the head
and prothorax of the fourth stage larva; B, lateral aspect of
the terminal abdominal segments of the fourth stage larva; C,
a dorsal aspect of the basimere, lateral aspect of the phallosome
of the male terminalia; D, lateral aspect of male antennal
flagellomeres V through X.
70 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
of the thorax separate the adult female from other members of the fraudatrix
eroup. The adult male may be recognized not only by the above mentioned
characters, but also by the presence of an inconspicuous tuft of narrow,
tapering, pointed scales on antennal flagellomere V. The fourth stage larva
is distinguished by abdominal hair 7-I single, thoracic hair 3-P with from 4
to 10 branches, and by the thoracic integument strongly spiculate.
FEMALE. Head. Decumbent scales of the vertex with a small central
patch of fine, pale brown scales, followed by a wide area of broader dark |
scales and large lateral areas of broad, pale scales; erect scales uniformly
dark brown. Thorax. Scutum with the integument pale brown with a distinctly
reddish tint, sparsely covered with dark scales; integument of pleuron similar
in color to the integument of the scutum, without a lower mesepimeral bristle.
Abdomen. Terga brown, with indistinct, palish anterior and posterior bands
on each tergum of some specimens; sterna pale.
MALE. Head. Proboscis with rather long dorsal setae and shorter,
forward projecting ventral hairs as well as stronger basoventral setae;
palpus with very small basal processes. Antenna. (Figure 20D). Flagello-
mere V with a tuft of from 2 to 5 long, dark, sharply pointed scales, the longest
of which extends to the apex of IX followed by approximately 7 long setae which
are stronger than those of the normal whorl]; VI with an internal tuft of 10 or
more short, stout, dark and twisted spines; VII with a similar, but smaller
tuft of spines; VIII with a dense tuft of long, gently curved spines; [X with from
3 to 5 well developed, lanceolate scales each of which terminates in a sharp
point; X with an internal tuft of from 2 to 4 strong, straight, dark setae.
Terminalia. (Figure 20C). Basimere with 3 long, strongly curved, sub-
marginal setae inserted in a straight row; subapical lobe of the basimere with
the internal rod expanded and sharply pointed apically, central and external
rods subequal in length and thickness and hooked apically; 6 accessory setae
present, all rather blade-like, and of variable length and with one of the
shorter setae serrate; internal and external leaflets subequal in length, sym-
metrical; distimere normal but with fine annulations on the convex surface at
the apex; dorsal process of the lateral plate of the phallosome long, strongly
curved, and sharply pointed, the posterior angle of which is smooth.
LARVA. (Figure 20A,B). Head. Antenna with a narrow, dark basal
band and darker beyond the insertion of hair 1-A; head hair 4-C with from 2
to 4 branches, simple; 5,6-C usually bifid, pectinate, occasionally single or
trifid on 1 side; 16,17-C represented by minute, but well developed spicules.
Thorax. Integument covered with a dense pattern of prominent, short spicules;
hair 3-P with from 4 to 10 pectinate branches, the individual branches consid-
erably shorter and slenderer than 1,2-P; 4-P bifid, pectinate; 5, 6-P single,
pectinate; 7-P with 3 or 4 branches, pectinate; 8-P bifid, pectinate; 14-P
single, simple. Abdomen. Integument glabrous; hair 7-I single; comb consist-
ing of from 35 to 50 scales arranged in a broad, triangular patch; siphon index
variable, ranging from 6:1 to 8:1 (average, 7.3:1); 4 pairs of subventral tufts
Figure 20. C. (Lophoceraomyia) rubithoracis. A, dorsoventral aspect of
the head and prothorax of the fourth stage larva; B, lateral
aspect of the terminal abdominal segments of the fourth stage
larva; C, dorsal aspect of the basimere, lateral aspect of the
phallosome of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through X.
—
Bram: Genus Culex in Thailand
TS
MENTE, gi
Uf, Yes ra xc hor
Ky | Wy
WY
C. (Lophoceraomyia) rubithoracis
T.L. HWANG
71
72 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
inserted in a line on the siphon; individual tufts with from 2 to 4 branches,
simple, their length approximately equal to the width of the siphon at the point
of insertion; pecten consisting of from 7 to 10 teeth restricted to the basal _
third or less of the siphon; individual pecten tooth with a fine distal spine and
from 8 to 12 subequal lateral barbs.
TYPE DATA. Lectotype male (terminalia mounted on a celluloid point)
from Kuala Lumpur in the British Museum.
DISTRIBUTION. In THAILAND, this species has been collected from:
Chanthaburi, Chiang Mai, Chon Buri, Krung Thep, and Ubon Thani. This.
species has also been recorded from SINGAPORE, MALAYA, INDIA, INDO-
NESIA, CHINA, BORNEO, JAPAN, TAIWAN, and the PHILIPPINES.
During this study 8 larvae were studied from Thailand as well as 50
males and 9 females; 1 male and 1 female with their associated larval and
pupal skins have been studied from Singapore.
TAXONOMIC DISCUSSION. As indicated in the above diagnosis, there
is little difficulty in distinguishing this species from other members of the
fraudatrix group. Specimens are occasionally found in which the characteris-
tic reddish brown tint of the thoracic integument is replaced by darker brown
color; in these cases it is necessary to resort to anatomical features before
drawing final conclusions.
BIOLOGY. Although the larvae have been infrequently collected in
Thailand, the adults were taken in rather large numbers at light traps near
Bangkok. The larvae are collected primarily in open areas such as ponds
and paddy fields; however, Colless (1965) found that even where larvae were
common, the adults were rarely, if ever, ‘ace on mammalian baits and
presumably fed on birds.
CULEX (LOPHOCERAOMYIA) VARIATUS (LEICESTER) 1908
(Figure 21)
Lophoceratomyia variata Leicester 1908, Cul. Malaya: 121 (“,?); Edwards
1913, Bull. ent. Res. 4: 234 (synonymy with fraudatrix).
Culex (Lophoceraomyia) variatus (Leicester): Colless 1965, J. med. Ent.
2: 270 (revalidated, **,?, L*).
The adult male is distinguished from other species of the fraudatrix
group by the presence of 3 submarginal setae in a line on the basimere, 8 or
more blunt scales on antennal flagellomere V, and by a Sharply angulate tuft
of scales on flagellomere VII. The fourth stage larva may be separated
from other species of the fraudatrix group by head hair 5-C being bifid and
long, abdominal hair 7-I branched, thoracic hair 4-P branched, and 14-P
single.
FEMALE. Thorax. One lower meSepimeral bristle present. Abdo-
men, Terga covered with dark brown scales, with small indistinct baso-
lateral pale patches on tergum VII; sterna with pale scales and often with in-
distinct subapical dark bands on the more posterior segments.
MALE. Head. Proboscis with rather long dorsal setae and shorter,
forward projecting ventral hairs, as well as stronger basoventral setae;
palpus with short basal processes, the outer, longer processes about half
the length of palpal segment II. Antenna. (Figure 21D). Flagellomere V
with a prominent external tuft of 8 or more broad, bluntly rounded scales
Bram: Genus Culex in Thailand 713
which extend to the tuft of X; VI with an internal tuft of 15 or more short,
stout, dark and twisted spines; VII with a similar, but smaller tuft of spines
and a secondary longer tuft; VIII with a tuft of long, broad spines which are
sharply angled at the apex; IX with from 4 to 6 finely pointed, lanceolate,
translucent leaflets as well as several stout setae; X with an internal tuft of
from 2 to 4 strong, dark setae. Terminalia. (Figure 21C). Basimere with
3 strong, slightly curved submarginal setae inserted in a straight row; sub-
apical lobe of the basimere with internal rod having an expanded apex, ex-
ternal and central rods subequal in length, hooked apically; 5 or 6 accessory
setae present, 2 of which are longer than the others, rather straight, and
blade-like; internal leaflet narrow basally, then expanded and blunt distally,
asymmetrical; external leaflet rather narrow, pointed apically; distimere
normal, but with prominent apical annulations on the convex margin; dorsal
process of the lateral plate of the phallosome long, gently curved, and sharp-
ly pointed, the posterior angle of which frequently exhibits several minute,
Spiculate teeth.
LARVA. (Figure 21A,B). Head, Hair 4-C simple, single or with up
to 4 branches; 5, 6-C bifid, pectinate, subequal in length; 16, 17-C represented
by minute spicules. Thorax. Integument covered with a rather dense pattern
of minute spicules; hair 3-P very fine, pectinate, considerably shorter than
1,2-P, either single or bifid; 4-P bifid, pectinate; 5,6-P single, pectinate;
7-P trifid, pectinate; 8-P bifid, pectinate; 14-P single, simple. Abdomen.
Integument glabrous; hair 7-I branched; 2-VIII bifid; comb consisting of from
35 to 55 fan-shaped scales arranged in a broad, triangular patch; siphon
index variable, ranging from 6. 5:1 to 9:1 (average, 7.5:1), a narrow median
dark band usually present; 4 pairs of subventral tufts inserted in a line on
the siphon (occasionally one side will exhibit either 3 or 5 tufts); individual
tufts with from 2 to 4 branches (occasionally single), simple, their length
equal to, or less than the width of the siphon at the point of insertion; pecten
consisting of from 9 to 13 teeth restricted to the basal fourth or less of the
- siphon; individual pecten tooth with a fine apical spine and from 6 to 11 fine
lateral barbs, a subbasal barb frequently more prominent than the other
lateral barbs.
TYPE DATA. Lectotype male (terminalia mounted and attached to
pin, antenna slide-mounted) from Kuala Lumpur, Malaya in the British Mu-
seum.
DISTRIBUTION. In THAILAND**, vaviatus has been studied from:
Krung Thep, Narathiwat, Ranong, Satun, Songkhla, and Thon Buri. Outside
of Thailand, the species has been recorded only from MALAYA.
During this study 9 males, 2 with their associated larval and pupal
skins, 3 females, 1 with associated skins, and 6 fourth stage larvae have
been examined.
TAXONOMIC DISCUSSION. This species demonstrates closest af-
finity to fraudatrix of New Guinea and, as pointed out by Colless (1965), has
probably been most frequently misidentified as that species. In the male,
variatus may be separated from fraudatrix on the basis of the lanceolate
leaflets on antennal flagellomere IX and the sharply angled spines on flagel-
lomere VIII; also, in fraudatrix the accessory setae on the subapical lobe of
the basimere are not as well developed as in varialus.
BIOLOGY. In Thailand, larvae have been collected from a variety of
eround water habitats including a marsh, a large ground pool, pools in dry
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
os
Ace:
1
ay
\
\
1
4
oem
C. (Lophoceraomyia) variatus
Bram: Genus Culex in Thailand 15
stream beds, anda sump, generally with floating leaves and other organic
matter. Colless (1965) suggested that the most common larval habitat of
this species is partly shaded and has much organic matter in the water,
which is stagnant and usually fresh; however, larvae were also collected
from container habitats near the ground such as palm axils, tins, cavities
in fallen logs or bamboos, and crab holes. Nothing is known of the adult
bionomics, but Colless (1965) has postulated that they may normally feed on
birds and forest mammals, and possibly act as vectors of certain viruses.
MAMMILIFER GROUP
FEMALE. Scales on wing veins Rg and Rg usually narrow, parti-
cularly on the basal half. Otherwise, consistent features are not presently
recognized which can be used to separate the groups within the subgenus.
MALE. Head. Torus of the antenna with a mammiliform protuber-
ance on the inner surface or (in the case of wilfredi), the torus asymmetrical
and with a slit-like depression on the inner surface; length of the palpus
variable; palpus without distinct basal processes; proboscis without long
dorsal setae, but usually with dorsal pubescence, particularly on the ex-
panded subbasal region; antenna usually with modified scales or setae on
flagellomeres V through IX. Terminalia. Subapical lobe of the basimere
usually with 1 of the leaflets modified; lateral plate of the phallosome with an
internal process which is smooth and gently curved, and a dorsal process
which is usually truncate with a toothed or spinosSe apex.
LARVA. Head. Length of 4-C distinctly greater than the distance
between the bases of the pair. Thorax. Hair 14-P bifid (except in
mammilifer and wildredi). Abdomen, Hair 2-VII single; siphon with 3 or
more pairs of subventral tufts; central filament of the siphon valves present
or absent, when present usually weakly developed.
DISTRIBUTION. Representatives of the mammilifer group are
restricted to the Oriental region. In Thailand, species of this group are un-
doubtedly distributed throughout the country.
TAXONOMIC DISCUSSION. Within the mammilifer group, 2 rather
well defined subgroups may be recognized in terms of anatomical features as
well as larval habitats. Further differentiation of these subgroups will be
given below.
BREVIPALPUS SUBGROUP
FEMALE. Anatomical features cannot presently be recognized which
separate members of the brevipalpus and mammilifer subgroups.
Figure 21. C. (Lophoceraomyia) variatus. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral
<— aspect of the terminal abdominal segments of the fourth stage
larva; C, dorsal aspect of the basimere, lateral aspect of the
phallosome of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through X.
76 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
MALE. Length of the palpus, at most, only slightly greater than that
of the proboscis (in the 2 species presently recognized from Thailand, the
palpus is only 1/2 to 3/4 the length of the proboscis). Distimere of the termi-
nalia noticeably expanded at the apex.
LARVA. Antenna cylindrical, not noticeably constricted beyond in-
sertion of hair 1-A; hairs 2-6-A all inserted at the apex of the shaft; mouth-
parts apparently adapted for predation.
LARVAL HABITAT. Normally, larvae of species of the brevipalpus
subgroup are found exclusively in pitcher plant (Nepenthes spp.) environments.
TAXONOMIC DISCUSSION. The short male palpus and the features of
the larval antenna clearly separate this subgroup from the mammilifer sub-
sroup. Of a total of at least 9 species which have been described from this
subgroup, only 2, curtipalpis and lucaris, have been positively identified
from Thailand. A single larva, collected from a pitcher plant in Ranong,
possibly represents brevipalpus (Theobald); however, the larvae of brevipalpus
and eminentia (Leicester) have not been adequately characterized to permit an
accurate determination.
CULEX (LOPHOCERAOMYIA) CURTIPALPIS (EDWARDS) 1914
(Figure 22)
Lophoceratomyia curtipalpis Edwards 1914, Bull. ent. Res. 5: 127 (¢).
Culex (Lophoceratomyia) curtipalpis (Edwards): Edwards 1921, Bull. ent.
Res. 12: 78 (as synonym of C. (Lophoceratomyia) jenseni (Meij. ));
Edwards 1928, Bull. ent. Res. 18: 279 (revalidated); Edwards and
Given 1928, Bull. ent. Res. 18: 357 (L*, P).
Culex (Lophoceraomyia) curtipalpis (Edwards): Colless 1965, J. med. Ent.
2° 301 (o*, ¢, 1). :
The adult female may be recognized by the absence of a lower mesep-
imeral bristle, by the presence of some Scales on the pleuron, and by the
presence of some pale, erect scales on the vertex. The adult male is
distinctive in the expanded apex of the distimere, the short palpus, and the
structures of the subapical lobe of the basimere. The short siphon of the
fourth stage larva distinguishes this species from all other known members
of the genus in Thailand.
FEMALE. Head. Decumbent scales of the vertex dark brown, but
with a relatively wide band of pale, broad scales adjacent to the orbital line;
erect scales dark brown with a golden tinge on the occiput and usually pale at
the lateral margins. Thovax. Integument of the pleuron usually dark brown,
sometimes paler, with rather distinctive patches of pale scales on the sterno-
pleuron and sometimes on the anterior pronotal lobe; lower mesepimeral
bristle absent. Legs. Anterior surface of hind femur predominantly pale,
with a narrow, dark apical band which extends proximally along the dorsal
margin; anterior surface of the mid femur similar to the hind femur, but with
the dark area considerably expanded. Abdomen. Terga dark scaled with a
paler stripe down each lateral margin; sterna pale.
MALE. Head. The length of the palpus at least 1/4 but less than 1/2
the length of the proboscis. Antenna. (Figure 22D). Flagellomere V witha
tuft of 2 or 3 long, strong, straight spines followed by 4 to 7 narrow, pointed,
darker scales; VI with a tuft of 4 or more simple, straight, rather short
Bram: Genus Culex in Thailand 17
spines; VII with a tuft of 5 short, hooked setae and from 3 to 5 longer,
straight setae; VII with a tuft of 3 or 4 narrow, lanceolate blades; IX witha
tuft of from 3 to 6 strong, straight setae. Terminalia. (Figure 22C). Basi-
mere without strongly distinctive submarginal setae; subapical lobe of the
basimere with the internal rod very strong, long, bluntly rounded and charac-
teristically curved, and with the central and external rods subequal in length,
shorter than the internal rod, and gently hooked; internal leaflet represented
by a stout, curved spine; external leaflet very broad, asymmetrical; 4 sharply
angled, blade-like, characteristically pointed accessory setae present; disti-
mere prominently expanded at the apex; dorsal process of the lateral plate of
the phallosome with a spiculate apical knob and with 15 or more fine, but
distinct, teeth on the lateral margin; internal process very short, gently
curved, and pointed.
LARVA. (Figure 22A,B). Head, Antennal shaft cylindrical, hair 1-A
very short, with from 2 to 4 simple branches, hairs 2-6-A inserted at the
apex of the shaft; head hair 1-C lightly pigmented, stout, very short, and
inserted in a large socket; 4-C simple, with from 1 to 3 branches, inserted
rather close to the anterior margin of the head; 5, 6-C fine, pectinate, with
from 2 to 4 branches, and also inserted well forward; 16,17-C absent;
ventral and internal mouth brushes very stout, hooked, apparently adapted
for predation. Thorax. Integument glabrous; 1-P expanded basally; 3-P
single, pectinate, shorter and slenderer than 1,2-P; 4, 7-P single or bifid,
pectinate; 5,6-P single, pectinate; 8-P extremely short, fine, simple, single
or bifid; 14-P single or bifid, simple. Abdomen. Integument glabrous; comb
consisting of an irregular row of from 4 to 10 blunt, elongated scales which
are fringed with fine, subequal spines; siphon shorter than the saddle on seg-
ment X, index ranging from 1. 2:1 to 1. 4:1; 3 or 4 pairs of subventral tufts
inserted in a line on the siphon; individual tufts with from 3 to 6 branches,
pectinate, the apical tuft extremely small and simple; pecten consisting of 3
or 4 stout, simple teeth restricted to the basal half of the siphon.
TYPE DATA. Holotype male (terminalia slide-mounted) from Kuching,
Sarawak in the British Museum.
DISTRIBUTION. In THAILAND**, specimens of this species have been
collected from Phatialung and Ranong. The species has also been reported
from SINGAPORE, SARAWAK, MALAYA (Colless 1965). Specimens of
curtipalpis have been examined in the U. S. National Museum collection from
Saigon (coll. H. H. Stage, 1955) and DaNang (coll. P. J. Santana, 1965),
SOUTH VIETNAM**.
The following specimens have been examined during this study: 4 fe-
males with their associated larval and pupal skins; 8 males, 4 with their
associated larval and pupal skins; and 76 larvae from South Vietnam.
BIOLOGY. Prior to this study, larvae of curtipalpis had been collected
exclusively from pitcher plants; however, on one occasion larvae of this
species were collected from a tree hole in Phattalung, Thailand. The larvae
apparently feed on insect remains which accumulate in the pitchers or tree
holes. In Thailand, collections have been made during the months of March
and October; in South Vietnam, during December, January, March, and May.
Habits of the adults are unknown.
78 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
JL. HWANG
Pees Ge eed
C.(Lophoceraomyia) curtipalpis
Bram: Genus Culex in Thailand 719
CULEX (LOPHOCERAOMYIA) LUCARIS COLLESS 1965
(Figure 23A, B)
Culex (Lophoceraomyia) lucaris Colless 1965, J. med. Ent. 2: 299 (c'*).
The adult male may be recognized by the palpus being from 1/2 to 3/4
the length of the proboscis, by the expanded apex of the distimere, and by
the characteristic tuft of scales on antennal flagellomere V.
FEMALE. Unknown.
MALE. Head. Proboscis and palpus dark brown, the length of the
palpus from 1/2 to 3/4 the length of the proboscis; erect scales of the vertex
pale bronze-brown on the occiput, becoming darker posterolaterally.
Antenna. (Figure 23B). Flagellomere V with a tuft of 6 or 7 golden, nar-
row, tapering scales whose apices extend beyond the tuft on IX; VI witha
tuft of about 4 rather long, gently sigmoid setae with filamentous apices and
5 or more Straight, slender setae; VII with a tuft of about 12 shorter scales
which terminate in a recurved point and a supplementary tuft of broader
scales; VIII with a tuft of 5 or more strong, gently curved setae; IX with
from 6 to 10 fine, straight setae. Thovax. Integument of the pleuron uni-
formly pale brown with one lower meSepimeral bristle present. Abdomen.
Terga covered with dark brown scales; sterna uniformly pale. Terminalia.
(Figure 23A). Basimere without strong, distinctive submarginal setae; sub-
apical lobe of the basimere with the internal rod stout, bent, shorter than the
central and external rods, and with a filamentous apex; central rod rather
narrow, smoothly curved apically; external rod longer than the other 2, ex-
panded and hooked at the apex; internal leaflet represented by a rather long,
rod-like structure; external leaflet not recognizable, possibly represented by
a moderately long seta; 5 accessory setae present, the 2 near the rods strong
and gently curved, the 3 between the rods and the leaflets short, rather broad,
and blade-like; a dense patch of short, fine setae also present around the
base of the leaflets; distimere with an expanded apex and exhibiting 3 strong
apical spines; dorsal process of the lateral plate of the phallosome with a
Spiculate apex and 7 or more strong teeth on the lateral margin; internal
process short, strongly curved and pointed.
LARVA. Unknown. :
TYPE DATA. Holotype male (terminalia and antenna slide- mounted)
from Singapore in the Australian National Insect Collection, Canberra.
DISTRIBUTION. C. lucaris is known only from the type locality (Ulu
Pandan, SINGAPORE) and from Ranong, THAILAND**.
The author has examined 1 paratype male (in the British Museum) and
a Single male from Thailand.
TAXONOMIC DISCUSSION. This species is extremely close to
eminentia (Leicester), but may be separated on the basis of the subapical
Figure 22. C. (Lophoceraomyia) curtipalpis. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
of the terminal abdominal segments of the fourth stage larva;
styl C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
80 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
|! as
Nest heise
Wye
LAD eee
C.(Lophoceraomyia) /ucaris
JL. HWANG
C. (Lophoceraomyia) incomptus
Bram: Genus Culex in Thailand 81
lobe of the basimere and the flagellomere tufts. In eminentia the subapical
lobe itself is rather narrow and conical, whereas this structure in lucaris
is broad and truncate; although the appendages on the subapical lobe of the 2
Species are very Similar, the accessory setae are relatively longer in
eminentia. :
BIOLOGY. The single specimen from Thailand was reared from a
pupa collected in a pitcher plant. Specimens of the type series were col-
lected resting in rather open forest.
MA MMILIFER SUBGROUP
MALE. Length of the palpus greater than that of the proboscis (ex-
cept in tvaubi, in which the palpus is never as short as 3/4 the length of the
proboscis); distimere of the terminalia not expanded apically.
LARVA. Antenna noticeably constricted beyond insertion of hair 1-A
which is plumose; antennal hairs 2,3-A inserted considerably below the apex
of the shaft, 4,5,6-A inserted at the apex; mouthparts not adapted for preda-
tion.
LARVAL HABITAT. Larvae are usually found in natural container
habitats such as tree holes, bamboo internodes, rock holes, etc., but are
also frequently found in a variety of ground water habitats.
TAXONOMIC DISCUSSION. Within the mammilifer subgroup, at
least one species complex is recognizable. This group includes tvaubi,
ganapathi, spiculosus, and peytoni, all of which exhibit a marked similarity
in anatomical features as well as larval habitats. Although these species
may be distinguished on the basis of the male terminalia and the tufts of the
male antennal flagellomeres, it is desirable to examine the associated larval
skins before drawing final conclusions.
CULEX (LOPHOCERAOMYIA) INCOMPTUS BRAM AND RATTANARITHIKUL
1967
(Figure 23C, D)
Culex (Lophoceraomyia) incomptus Bram and Rattanarithikul 1967, Proc. ent.
Soc. Wash. 69: 16 (<*).
The adult male may be recognized by the presence of 2 irregular rows
of submarginal setae on the basimere of the terminalia, by the 6 acutely
pointed scales on antennal flagellomere V, and by the characteristically bulbous
setae on flagellomere VII.
Figure 23. C. (Lophoceraomyia) lucaris. A, dorsal aspect of the basimere,
lateral aspect of the phallosome of the male terminalia; B,
lateral aspect of male antennal flagellomeres V through IX.
<< C. (Lophoceraomyia) incomptus. C, dorsal aspect of the basi-
mere, lateral aspect of the phallosome of the male terminalia;
D, lateral aspect of male antennal flagellomeres V through IX.
82 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
FEMALE. Unknown.
MALE. Antenna. (Figure 23D). Flagellomere V with an internal
tuft of 6 acute setae which extend to the tuft of VI; VI with a tuft of 6 rather
fine, short setae which are bent distally; VI. with a similar tuft of from 10
to 15 shorter setae; VII with an internal tuft of 7 strong, dark, gently curved
setae, the distal 2 of these setae with a characteristic bulbous, median ex-
pansion; IX with an internal tuft of 3 short, slender setae and 3 longer,
slenderer setae. Thorax. Integument of the pleuron uniformly pale, tinged
with green in Some specimens; 1 lower mesepimeral bristle present.
Abdomen. Terga dark brown; sterna slightly lighter. Terminalia. (Figure
23C). Basimere with from 9 to 11 strong, submarginal setae inserted in 2
irregular rows; subapical lobe of the basimere with the internal rod rather
robust, constricted at the apex and pointed, the central and external rods
subequal in length, gently curved; internal leaflet rod-like, external leaflet
broad, striate; accessory processes narrow and Setae-like; distimere with
minute annulations on the apical third of the convex surface; dorsal process of
the lateral plate of the phallosome with an apical spiculate knob and 12 or more
fine, but distinct, denticles on the lateral margin; internal process robust,
distinctly curved, not projecting beyond the apex of the dorsal process.
LARVA. Unknown.
TYPE DATA. Holotype male (terminalia and antennae slide-mounted)
from Doi Suthep, Chiang Mai, Thailand in the U. S. National Museum.
DISTRIBUTION. In addition to the type locality, this species is known
from Doi Tad Fah, Chiang Mai, THAILAND. During this study 4 adult males
and their associated slide mounted terminalia and antennae were examined.
TAXONOMIC DISCUSSION. This species demonstrates its closest
affinity to fuscosiphonis and demissus Colless (the latter presently known
only from Selangor, Malaya), but the bulbous expansion of setae on flagello-
mere VIII and the shape of the processes on the subapical lobe of the basi-
mere clearly separates incomptus from its close relatives. C. bandoengensis
Brug (another member of the mammilifer subgroup whose distribution is
restricted to Malaya and Java) also exhibits the submarginal setae in 2 rows
but may be easily separated on the basis of the shape and number of the ac-
cessory processes on the subapical lobe of the basimere.
- BIOLOGY. Larvae were collected (and the skins subsequently lost)
from a tree hole on Doi Suthep Mountain during the month of January; 1 adult
male was collected from a low plant in a shady damp jungle on Doi Tad Fah
Mountain during May. No other biological data are available.
CULEX (LOPHOCERAOMYIA) BENGALENSIS BARRAUD 1934
(Figure 24)
Culex (Lophoceratomyia) minor var. bengalensis Barraud 1934, Fauna Brit.
India, Diptera 5: 371 (c).
Culex (Lophoceraomyia) bengalensis Barraud: Colless 1965, J. med. Ent. 2:
293 (o*, 2, L*).
Except for the absence of a lower mesepimeral bristle, the adult female
cannot be distinguished from other members of the mammilifer subgroup.
The adult male may be recognized by the presence of a tuft of from 1 to 4 very
Bram: Genus Culex in Thailand 83
long, filamentous scales on flagellomere V, by the tuft of obviously modified
dark, strong setae on flagellomere IX, by the submarginal setae of the basi-
mere inserted in a single row, and by the absence of a lower mesepimeral
bristle. The fourth stage larva possesses only 3 pairs of subventral tufts,
thoracic hair 14-P bifid, and abdominal hair 6-III usually trifid.
FEMALE. Thovax. Integument of pleuron dull brown, without
distinct patches of pale scales; lower mesSepimeral bristle absent. Abdomen.
Terga dark; sterna somewhat paler.
MALE. Head. Proboscis with a series of 8 or more short, baso-
ventral bristles. Antenna. (Figure 24D). Flagellomere V with a tuft of from
1 to 4 long, strong setae which extend beyond the base of VIII; VI with an in-
ternal tuft of 6 or more broad, gently sigmoid setae which have filamentous
apices; VI with a dense tuft of 10 or more fine spines which are sharply bent
and a tuft of approximately 5 or more strong, darkly pigmented, gently
curved setae; VIII with a tuft of from 5 to 8 long, gently curved spines; IX
with a tuft of from 4 to 7 strong, straight setae. Terminalia. (Figure 24C).
Basimere with 4 or 5 submarginal setae inserted in a line; subapical lobe
with the internal rod expanded distally, frequently pointed, and slightly longer
than the central and external rods which are subequal in length and hooked;
internal leaflet bladelike, gently curved, external leaflet rather narrow,
symmetrical and striated; 4 narrow, subequal, bladelike accessory setae
present, at least 2 of which are hooked; distimere normal in shape, without
annulations or retrorse spines on the convex surface; lateral plate of the
phallosome with the internal process rather broad, gently curved and pointed,
not exceeding the apex of the dorsal process (not illustrated); dorsal process
with a spiculate apical knob and approximately 10 teeth on the lateral margin.
LARVA. (Figure 24A,B). Head. Antenna with a narrow dark basal
ring; head hair 4-C single, simple; 5, 6-C bifid, pectinate, subequal in length;
16,17-C developed as relatively long, simple spines. Thorax. Integument
glabrous; 3-P single, pectinate, shorter and slenderer than 1,2-P; 4-P bifid,
pectinate; 5,6-P single, pectinate; 7, 8-P bifid, pectinate; 14-P bifid, simple.
Abdomen. Integument glabrous; hair 6-II usually trifid; comb consisting of
from 35 to 50 fan-shaped scales arranged in a broad, triangular patch; siphon
index variable, ranging from 6.5:1 to 9:1 (average, 7.6:1); 3 pairs of sub-
ventral tufts inserted in a line on the siphon; individual tufts bifid, their length
greater than the width of the siphon at the point of insertion; pecten consisting
of from 12 to 15 teeth restricted to the basal third to fourth of the siphon; in-
dividual pecten tooth with a prominent apical spine and approximately 7 lateral
barbs which become progressively larger from apex to base.
TYPE DATA. Lectotype male (head and terminalia mounted on sep-
arate slides, the rest of the specimen missing) from Nongpoh, Assam, in
the British Museum.
DISTRIBUTION. In THAILAND**, this species has been collected
from Ranong and Narathiwat. The species is also known from ASSAM,
HAINAN ISLAND, and MALAYA. During this study the following specimens
of bengalensis have been examined: 6 males, 3 females, and 6 larvae.
TAXONOMIC DISCUSSION. In the adult stage this species demonstrates
its closest affinity to minor; however, the tuft on antennal flagellomere V and
the chaetotaxy of the basimere are distinctive. The fourth stage larva is
most similar to mammilifer and some specimens cannot always be separated
with certainty; differences in chaetotaxy, as summarized in the key, may be
useful in separating the 2 species.
84
Mt
ry
Y
AA i}
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
mee
er:
1.0mm
os]
0
JoL . HWANG
|
C.(Lophoceraomyia) bengalensis
Bram: Genus Culex in Thailand 85
BIOLOGY. In Malaya, larvae were collected from a rock pool, a
streamside pool, anda tree hole. In Thailand, collections have been made
from elephant hoof prints, a stream margin, and a tree stump hole during
the month of September. Nothing is known of the adult habits.
CULEX (LOPHOCERAOMYIA) EUKRINES BRAM AND RATTANARITHIKUL 1967
(Figure 25) :
Culex (Lophoceraomyia) eukrines Bram and Rattanarithikul 1967, Proc. ent.
Soc. Wash. 69: 11 (o*, 2, L*).
The adult male exhibits an internal tuft with from 3 to 5 gently curved
setae on antennal flagellomere VII and a tuft of obviously modified, dark,
strong setae on flagellomere IX. The fourth stage larva may be immediately
recognized by the head hair 6-C which is always single and stout.
FEMALE. Thorax. Integument of the pleuron pale, without distinct
patches of scales; 1 lower mesepimeral bristle present. Abdomen. Terga
dark brown; sterna somewhat lighter.
MALE. Head, Proboscis with a series of 6 or more short, baso-
ventral bristles. Antenna. (Figure 25D). Flagellomere V with a tuft of 6
or more very fine setae extending beyond the tuft of IX; VI with a tuft of 5 or
more gently curved setae; VII with an internal tuft of from 3 to 5 gently sig-
moid setae and several shorter, straight setae; VIII with a tuft of from 4 to 6
strong, gently curved setae; IX with a tuft of from 3 to 5 long, strong, dark
setae. Terminalia. (Figure 25C). Basimere with 5 or 6 rather weak sub-
marginal setae; subapical lobe of the basimere with the internal rod slightly
expanded subapically, then bent and with an extended, lightly sclerotized
hooked apex; central and external rods subequal in length, sharply hooked;
4 blade-like accessory processes present; internal leaflet rod-like, gently
bent; external leaflet slender, striate; distimere normal, without distal ser-
rations or annulations; lateral plate of the phallosome with the internal
process rather small, gently curved, not exceeding the apex of the dorsal
process; dorsal process with an apical spiculate knob and approximately 15
short, but distinct teeth on the lateral margin.
LARVA. (Figure 25A,B). Head. Antenna with a narrow, dark basal
ring and progressively darker beyond insertion of hair 1-A; head hair 4-C
single, simple; 5-C usually bifid, sometimes single, pectinate; 6-C always
single, pectinate, somewhat stouter than the individual branches of 5-C;
16,17-C represented by minute spicules. Thorax. Integument glabrous; 3-P
single, pectinate, shorter and slenderer than 1,2-P; 4-P bifid, pectinate;
5,6-P single, pectinate; 7-P single or bifid, pectinate; 8-P single, pectinate;
14-P bifid, simple. Abdomen. Integument glabrous; comb consisting of from
Figure 24. C. (Lophoceraomyia) bengalensis. A, dorsoventral aspect of
the head and prothorax of the fourth stage larva; B, lateral
, aspect of the terminal abdominal segments of the fourth stage
<— larva; C, dorsal aspect of the basimere and distimere of the
male terminalia; D, lateral aspect of male antennal flagello-
meres V through IX.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
!
4
A
4 bh
He
Ky
C.(Lophoceraomyia) eukrines
—T-L. HWANG
Bram: Genus Culex in Thailand 87
35 to 50 elongate, fan-shaped scales arranged in a broad triangular patch;
siphon index variable, ranging from 7:1 to 10:1 (average, 8.2:1); 4 pairs of
subventral tufts inserted in a line on the siphon; individual tufts with from 2
to 4 branches, their length greater than the width of the siphon at the point
of insertion; pecten consisting of from 8 to 12 teeth restricted to the basal
third to fourth of the siphon; individual pecten tooth with a prominent distal
Spine and from 5 to 9 lateral barbs, the proximal 2 or 3 barbs more promi-
nent than the distal barbs.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia slide-mounted) from Banchanunt Thailand in the U. S. National
Museum.
DISTRIBUTION. This species is known from 24 collections from
Kanchanaburi and 1 collection each from Nakhon Nayok and Trang, THAI-
LAND.
During this study 14 individual rearings were examined, as well as 17
additional males, 21 additional females, and 120 larvae.
TAXONOMIC DISCUSSION. The adult stage (both male and female)
demonstrates affinity to the traubi-ganapathi species complex but the distinct-
ive larva seems to exclude it from this group. The adult male is differentiated
from the member of the ‘vaubi-ganapathi complex by the presence of an in-
ternal tuft of from 3 to 5 gently sigmoid setae on flagellomere VI; it differs
from peytoni by lacking accessory setae on the basal third of the distimere and
by possessing approximately 15 teeth on the lateral margin of the dorsal
process of the phallosome; it differs from spiculosus on the basis of the in-
ternal rod on the subapical lobe of the basimere. The fourth stage larva is
recognized by head hair 6-C which is always single, stout, and apparently
somewhat flattened.
BIOLOGY. Larvae of eukrines have been collected predominantly from
bamboo internodes, but additional collections have been made from rock holes,
flood pools, a coconut shell, a wheel track, a tree hole, and a pandanus axil,
always in close association with bamboo groves in rain forest. Collections
were made during the months of March, and May through October. Nothing is
known of the habits or biology of the adults.
CULEX (LOPHOCERAOMYIA) FUSCOSIPHONIS BRAM AND RATTANARITHIKUL 1967
(Figure 26)
Culex (Lophoceraomyia) fuscosiphonis Bram and Rattanarithikul 1967, Proc.
ent. Soc. Wash. 69: 14 (o*, 9, L*).
The adult male is recognized by flagellomere V lacking a conspicuous
tuft of long, broad scales, by having the prominent submarginal setae inserted
Figure 25. C. (Lophoceraomyia) eukrines. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
aes
88
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967—
cs
a
14
Vill
5 WshAliad
LO ales,
Mi pane Py 3 2 4
ren hh
“ARS nNsa,
a? VA
C. (Lophoceraomyia) fuscosiphonis Jr. HWANG
Bram: Genus Culex in Thailand 89
in 2 irregular rows on the basimere, and by the internal process of the
lateral plate of the phallosome projecting beyond the apex of the dorsal
process. The fourth stage larva exhibits a long, darkly pigmented siphon
with 4 pairs of fine, rather short subventral tufts and thoracic hair 8-P is
single, simple.
FEMALE. Thorax. Integument of the pleuron pale, without distinct
patches of scales; 1 lower mesepimeral bristle present. Abdomen. Terga
dark brown; sterna somewhat lighter.
MALE. Head. Proboscis with a series of 6 or more short, baso-
ventral bristles. Antenna. (Figure 26D). Flagellomere V with an internal
tuft of from 5 to 7 narrow, acute setae whose apices do not reach beyond VII;
VI with a tuft of 6 or more strong, bent setae; VII with a similar, but shorter
and more prominently sigmoid tuft of setae; VIII with a tuft of approximately
6 strong, gently curved setae; IX with a tuft of from 3 to 5 strong, rather
long, dark setae. Terminalia. (Figure 26C). Basimere with approximately
9 submarginal setae inserted in 2 parallel, but irregular, rows; subapical
lobe of the basimere with the internal rod rather broad, constricted at the
apex and filamentous beyond the constriction; the central and external rods
subequal in length, hooked; internal leaflet rod-like; external leaflet slender,
striate; 4 short, accessory setae present; distimere normal, without apical
annulations; lateral plate of the phallosome with the internal process rather
narrow, gently curved and pointed, projecting beyond the apex of the dorsal
process; dorsal process with a spiculate apical knob and approximately 10
short teeth on the lateral margin.
LARVA. (Figure 26A,B). Head. Antenna with a narrow, dark basal
ring and darker beyond insertion of 1-A; head hair 4-C single, simple; 5, 6-C
bifid, pectinate; 16,17-C absent. Thorax. Integument covered with a sparse
pattern of extremely fine, rather long spicules; 3-P single, pectinate, shorter
and Slenderer than 1,2-P; 4-P bifid, pectinate; 5,6-P single, pectinate; 7-P
bifid, pectinate; 8-P single, simple; 14-P bifid, simple. Abdomen. Integu-
ment covered with a pattern of spicules similar to that of the thorax; hair 6-I
trifid; comb consisting of from 30 to 45 elongate, fan-shaped scales arranged
in a broad triangular patch; siphon index variable, ranging from 10:1 to 12. 6:1
(average, 11. 8:1); 4 pairs of subventral tufts inserted in a line on the siphon;
individual tufts fine, with from 2 to 4 branches, their length approximately
equal to the width of the siphon at the point of insertion; pecten consisting of
from 7 to 10 teeth restricted to the basal fourth or less of the siphon; indi-
vidual pecten tooth slightly curved, with a prominent distal spine and with from
6 to 10 lateral barbs; distal barbs fine, proximal barbs coarse and conspic-
uous; siphon and anal segment very darkly pigmented, more so than in any
other species of the subgroup. The association of the larva with the male is
presumptive; larvae were collected along with a pupa from which the holotype
male was reared.
Figure 26. C. (Lophoceraomyia) fuscosiphonis. A, dorsoventral aspect of
the head and prothorax of the fourth stage larva; B, lateral
as aspect of the terminal abdominal segments of the fourth stage
larva; C, dorsal aspect of the basimere, lateral aspect of the
phallosome of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
90 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
TYPE DATA. Holotype male (terminalia and antennae slide-mounted)
from Phattalung, Thailand in the U. S. National Museum.
DISTRIBUTION. This species is known from THAILAND from:
Phattalung, Trat, and Trang. During this study 2 males reared from pupae
have been examined, as well as 1 female with associated larval and pupal
skins, and 13 larvae.
TAXONOMIC DISCUSSION. The adult male of fuwscosiphonis demon-
strates close affinity to demissus (a species known only from Selangor,
Malaya) and cannot be separated on the basis of the terminalia alone; how-
ever, the internal tuft of antennal flagellomere V in fuscosiphonis does not
exhibit the distinctively long setae which are said to be present in demissus.
The fourth stage larva exhibits close similarities with wilfredi, but may be
distinguished on the basis of the very darkly pigmented siphon and anal seg-
ment and by hair 2 of the anal segment which is bifid in the case of
fuscosiphonis and trifid in wilfredi.
BIOLOGY. Larvae have been collected in tree holes and root holes
in a primary rain forest and a secondary scrub stand. Nothing is known of
the adult habits or biology.
CULEX (LOPHOCERAOMYIA) GANAPATHI COLLESS 1965
(Figure 27)
Culex (Lophoceraomyia) ganapathi Colless 1965, J. med. Ent. 2: 294 (c*, 9,
L*),
The adult male may be recognized by the single row of submarginal
setae on the basimere of the terminalia, by the absence of a tuft of modified
setae on antennal flagellomere IX, and by the tuft of long, narrow, pointed
scales on flagellomere V. The fourth stage larva possesses 4 pairs of
strong, subventral tufts on the siphon, the thoracic integument covered with
fine spicules, the abdomen glabrous, and head hair 1-C usually expanded on
the basal half.
FEMALE. Thorax. Integument of the pleuron dull brown, without
distinct patches of pale scales; 1 lower mesepimeral bristle present. Abdo-
men. Terga dark; sterna somewhat paler.
MALE. Head. Proboscis without distinctive dorsal or ventrolateral
setae, but with a series of 6 or more short, basoventral bristles. Antenna.
(Figure 27D). Flagellomere V with a tuft of from 3 to 6 narrow, tapering,
pointed scales which almost extend to the tuft on VIII; VI with a tuft of 10 or
more strong, gently curved scales which have filamentous apices; VII witha
tuft of 7 or more narrow, rather short, sharply bent scales followed by from
3 to 6 stronger, longer and kinked scales; VIII with a prominent tuft of 6 or
more strong, darkly pigmented curved setae; [IX without modified scales or
Figure 27. C. (Lophoceraomyia) ganapathi. A, dorsoventral aspect of
the head and prothorax of the fourth stage larva; B, lateral
aspect of the terminal abdominal segments of the fourth stage
larva; C, dorsal aspect of the basimere, lateral aspect of the
phallosome of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
pe
Bram: Genus Culex in Thailand
C.(Lophoceraomyia) ganapathi
91
92 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
setae. Terminalia. (Figure 27C). Basimere with from 4 to 6 fine sub-
marginal setae inserted in a line; subapical lobe of the basimere with the
internal rod slightly expanded subapically, then tapering to a fine hook;
central and external rods close to and slightly longer than the internal rod,
hooked; internal leaflet strong, rod-like, expanded and gently bent on the
central third; external leaflet rather short, narrow, symmetrical, slightly
pointed apically and striate; 3 rather broad, subequal, curved accessory
setae present; distimere normal in shape, occasionally with a fine, indistinct
crest on the apex (not illustrated); lateral plate of the phallosome with the
internal process broad, gently curved and pointed; dorsal process with an
apical knob which is spinose only on the apical surface and with approximately
15 denticles on the lateral margin.
LARVA. (Figure 27A,B). Head. Antenna with a narrow, dark basal
ring and darker beyond insertion of 1-A; head hair 1-C darkly pigmented,
usually swollen on the basal half, then abruptly narrowed or occasionally
branched; 4-C single (infrequently bifid on 1 side), very finely and sparsely
barbed (a condition too fine to be shown in the illustration); 5, 6-C bifid,
pectinate, subequal in length; 16,17-C represented by extremely minute,
rather closely placed spines which may be bifid. Thorax. Integument
covered with fine spicules; 3-P single, pectinate, shorter and slenderer than
1,2-P; 4-P bifid, pectinate; 5,6-P single, pectinate; 7-P bifid, pectinate;
8-P single, pectinate; 14-P usually bifid, simple (occasionally single or
trifid on one side). Abdomen. Integument mostly smooth; hair 6-I trifid;
comb consisting of from 35 to 45 fan-shaped scales arranged in a broad, tri-
angular patch; siphon index variable, ranging from 7:1 to 11:1 (average,
8.3:1); 4 pairs of subventral tufts inserted in a line on the siphon; individual
tufts with from 3 to 5 strong branches of decreasing length from base to apex,
the length of the basal pair approximately 4 times the width of the siphon at
the point of insertion; pecten consisting of from 9 to 14 teeth restricted to
less than the basal third of the siphon; individual pecten tooth with a fine
distal spine and from 5 to 9 graded lateral barbs.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia slide-mounted) from Selangor, Malaya in the Australian National
Insect Collection, Canberra.
DISTRIBUTION. Specimens of sanapathi have been studied from 123
collections in THAILAND** from: Chanthaburi, Chiang Mai, Chumphon,
Kampong Pate, Kanchanaburi, Nakhon Nayok, Nakhon Ratchasima, Narathiwat,
Phattalung, Ranong, Tak, and Trang. This species is also known from Ulu
Langat, Ulu Gombak, and Rantau Panjang, MALAYA.
The following specimens were examined during the course of this
study: 61 males, 38 of which had associated larval and pupal skins; 29 females
with their associated larval and pupal skins; and 308 larvae. Included among
the specimens examined were male and female paratypes, both with associated
larval and pupal skins.
TAXONOMIC DISCUSSION. C. ganapathi belongs to the species complex
which also includes traubi, spiculosus, and peytoni. These 4 species are in-
distinguishable as females and the males may be recognized primarily on the
basis of the specialized scales and setae on flagellomeres V through IX; the
fourth stage larvae are also similar, but differences exist in integumental
Spiculation and chaetotaxy.
BIOLOGY. In Malaya this species was collected primarily from larval
Bram: Genus Culex in Thailand 93
habitats in association with bamboo and secondarily from tree holes. In
Thailand, 52 collections were made from tree holes and 44 from bamboo,
suggesting an equal preference for these two types of habitats. Other habitats
from which larvae were collected included: root holes and stump holes,
stream pools, stream margins, rock pools, a coconut shell, a can, a water
jug, and a pandanus axil. Collections were made from April through Novem-
ber. Nothing is known of the adult biology.
CULEX (LOPHOCERAOMYIA) MAMMILIFER (LEICESTER) 1908
(Figure 28)
Lophoceratomyia mammilifer Leicester 1908, Cul. Malaya: 128 (c, 2);
Edwards 1913, Bull. ent. Res. 4: 236 (placed bicornutus as synonym).
Culex (Lophoceratomyia) mammilifer (Leicester): Edwards 1922, Indian J.
med. Res. 10: 281, 473 (key); Barraud 1924, Indian J. med. Res. 12:
43 (o); Edwards 1928, Bull. ent. Res. 18: 276 (L); Barraud 1934,
Fauna Brit. India, Diptera 5: 374 (o'*, L); Baisas 1935, Philipp. J.
Sci. 57: 174 (o*).
Culex (Lophoceraomyia) mammilifer (Leicester): Bohart 1945, Navmed 580:
75 (distribution); Mattingly 1949, Proc. R. ent. Soc. Lond. (B) 18:
227 (L); Colless 1965, J. med. Ent. 2: 287 (o*, 9, L*); Delfinado 1966,
Mem. Amer. ent. Inst. 7: 109 (“*, L*, P*).
Culex (Lophoceratomyia) chiungchungensis Hsu 1963, Acta ent. Sinica 12:
231 («*, L*); Colless 1965, J. med. Ent. 2: 287 (synonymy).
The adult male may be recognized by the long, broad scales on
antennal flagellomere V and by the 6 to 8 submarginal setae on the basimere.
The fourth stage larva exhibits only 3 pairs of subventral tufts on the siphon
and thoracic hair 14-P is single.
FEMALE. Thorax. Integument of the pleuron dull brown, without
distinct patches of pale scales; 1 weak lower mesepimeral bristle present.
Abdomen. Terga uniformly dark brown; sterna pale.
MALE. Head. Proboscis without conspicuous dorsal setae, but with
numerous short ventral setae and a group of 10 or more short, basoventral
bristles. Antenna. (Figure 28D). Flagellomere V with a tuft of from 1 to 4
broad scales which extend to the tuft on VII, followed by 3 or 4 more ventral
scales which are sharply pointed and shorter, and several long hairs; VI with
a comb- like tuft of 10 or more short, dark, bent scales; VII with a similar
but shorter tuft in which the scales are more prominently bent and with a
secondary tuft of broader, longer scales; VIII with a tuft of 8 or more strong,
darkly pigmented and strongly curved setae; IX with 7 or more strong,
straight, modified hairs. Terminalia. (Figure 28C). Basimere with from 6
to 8 long, prominent submarginal setae which are distinctly bent medially;
subapical lobe of the basimere with the basal rods subequal in length, the
internal rod slightly expanded and curved distally; internal leaflet developed
as a rod-like structure, external leaflet broader, symmetrical, and minutely
striate; 4 or 5 accessory setae present; distimere normal in shape, with very
fine retrorse spines forming a crest on the convex surface of the apical fifth;
lateral plate of the phallosome with the internal process rather broad, gently,
but slightly curved, bluntly rounded and projecting beyond the apex of the
94
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C.(Lophoceraomyia) mammili fer
Bram: Genus Culex in Thailand 95
dorsal process; dorsal process with a spiculate apical knob and with from 3
to 6 small denticles on the lateral margin.
LARVA. (Figure 28A,B). Head. Antenna with a narrow, dark basal
ring and slightly darker beyond insertion of hair 1-A; head hair 4-C single,
simple, very long; 5, 6-C bifid, pectinate, subequal in length; 16,17-C
represented by well developed spicules which may be single but usually at
least 1 of the pair has up to 4 branches. Thorax. Integument usually glabrous,
but occasionally the prothorax exhibits light spiculation anterolaterally (not
illustrated); 3-P single, pectinate, shorter and slenderer than 1,2-P; 4-P
bifid, pectinate; 5,6-P single, pectinate; 7,8-P bifid, pectinate; 14-P single,
simple. Abdomen. Integument glabrous; hair 6-III with 4 branches; comb
consisting of from 33 to 45 fan-shaped scales arranged in a broad, triangular
patch; siphon index variable, ranging from 7. 5:1 to 12:1 (average, 9:1); 3
pairs of subventral tufts inserted in a line on the siphon; individual tufts
usually bifid, occasionally single, their length approximately equal to the
width of the siphon at the point of insertion; pecten consisting of from 10 to
14 teeth restricted to the basal third or less of the siphon; individual pecten
tooth with a fine distal spine and from 6 to 9 graded lateral barbs.
TYPE DATA. Lectotype male (terminalia and antenna slide-mounted)
of mammilifer from Raub, Pahang, Malaya in the British Museum. The type
locality of chiungchungensis is Chiungchung siang, Hainan Island, but the
location of the type specimen is unknown.
DISTRIBUTION. In THAILAND, mammilifer has been collected from:
Chiang Mai, Kanchanaburi, Lampang, Nakhon Ratchasima, Narathiwat, and
Yala. This species has also been recorded from MALAYA, INDIA, CEYLON,
ANDAMAN ISLANDS, HAINAN ISLAND, BORNEO, and the PHILIPPINES.
During this study 10 males (3 with associated larval and pupal skins)
have been studied in addition to 3 females with associated larval and pupal
skins, and 12 larvae.
TAXONOMIC DISCUSSION. This species demonstrates its closest
affinity to pholeter, since in the male both species exhibit a conspicuous tuft
of from 1 to 4 long, broad scales on antennal flagellomere V; however,
differences in the male terminalia of the 2 species are distinctive. In the
fourth stage larva, mammilifer, bengalensis, and minor possess only 3 pairs
of subventral tufts on the siphon, but they are readily separated by differences
in thoracic and abdominal chaetotaxy and in the latter species by the 2 distinct
types of comb scales.
BIOLOGY. Larvae have been commonly collected from container
habitats, axils of Nipa palms, and ground pools. Additional habitats from
Thailand include hoof prints, elephant tracks, seepage springs, and bamboo
internodes and stump holes. Collections were made during the months of
April, May, June, September, and October. Habits of the adults are unknown,
but they rarely, if ever, bite man and have, on occasion, been collected in
Figure 28. C. (Lophoceraomyia) mammilifer. A, dorsoventral aspect of
the head and prothorax of the fourth stage larva; B, lateral
aspect of the terminal abdominal segments of the fourth stage
<— larva; C, dorsal aspect of the basimere, lateral aspect of the
phallosome of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
96 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
light traps.
CULEX (LOPHOCERAOMYIA) MINOR (LEICESTER) 1908
(Figure 29)
Lophoceratomyia minor Leicester 1908, Cul. Malaya: 126 (<); Edwards
1917, Bull. ent. Res. 17: 227 (“, L); Borel 1926, Arch. Insts.
Pasteur Indo-Chine 3-4: 35 (o*, 9, L*); Borel 1930, Monogr. Coll.
Soc. Pat. exot. 3: 356 (o"*, 2, L*).
Lophoceratomyia bicornuta Theobald 1910, Rec. Indian Mus. 4: 25 (c);
Edwards 1932, in Wytsman, Genera Insect., fasc. 194: 198 (as
synonym of mammilifer).
Culex (Lophoceraomyia) bicornutus (Theobald): Colless 1965, J. med. Ent.
2: 291 (resurrected from synonymy with mammilifer, **,?, L*).
NEW SYNONYMY.
Culex (Lophoceratomyia) minor (Leicester): Edwards 1922, Indian J. med.
Res. 10: 282 (key); Barraud 1924, Indian J. med. Res. 12: 44 (o'*);
Edwards 1928, Bull. ent. Res. 18: 276 (key); Edwards and Given
1928, Bull. ent. Res. 18: 352 (L); Barraud 1934, Fauna Brit. India,
Diptera 5: 370 (o*, L); Safyanova et al. 1964, Zool. Zhur. 43: 1178
(distribution).
Culex (Lophoceratomyia) nolledoi Baisas 1935, Philipp. J. Sci. 57: 170 (o*);
Bohart 1945, Navmed 580: 76 (c‘*); Colless 1965, J. med. Ent. 2:
289 (Synonymy).
Culex (Lophoceraomyia) minor (Leicester): Mattingly 1949, Proc. R. ent.
Soc. Lond. (B) 18: 227 (key); Colless 1965, J. med. Ent. 2: 289 (o*,
2, L*); Delfinado 1966, Mem. Amer. ent. Inst. 7; 111 (o*, 9).
The adult male exhibits a tuft of obviously modified dark, strong »
setae on flagellomere IX, an internal tuft of at least 5 strong, sigmoid setae
on flagellomere VII, flagellomere V with a tuft of short, strong setae, 1
lower mesepimeral bristle, and lacks long, ventrolateral setae on the basal
half of the proboscis. The fourth stage larva possesses a comb with 2
distinct types of scales.
FEMALE. Thorax. Integument of the pleuron dull brown, without
distinct patches of pale scales; 1 lower mesepimeral bristle present. Abdo-
men. Terga dark; sterna somewhat paler.
MALE. Head. Proboscis without distinctive dorsal or ventrolateral
setae, but with a series of 8 or more short, basoventral bristles. Antenna.
(Figure 29D). Flagellomere V with a tuft of from 3 to 5 narrow, pointed
strong scales which extend only slightly beyond the tuft on VI and several
additional fine setae; VI with a comblike tuft of 10 or more dark, bent scales;
Figure 29. C. (Lophoceraomyia) minor. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; A', enlarged
pecten tooth of minor form bicornutus; B, lateral aspect of
the terminal abdominal segments of the fourth stage larva; C,
dorsal aspect of the basimere, lateral aspect of the phallosome
of the male terminalia; D, lateral aspect of male antennal
flagellomeres V through IX. ,
Bram: Genus Culex in Thailand
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98 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
VII with a similar but shorter tuft of 5 or more sigmoid setae and a secondary
tuft of broader, longer, sigmoid scales; VIII with a prominent tuft of 7 or
more strong, darkly pigmented, gently curved setae; IX with 5 or more strong,
straight setae. Terminalia. (Figure 29C). Basimere with from 5 to 7 sub-
marginal setae inserted in a straight line, the setae strong and prominently
curved; subapical lobe of the basimere with the internal rod separated from
the central and external rods, slightly expanded apically, then bent and
becoming narrow and terminating in a small hook; central and external rods
Subequal in length and strength, gently hooked; internal leaflet developed as
a rather long, rod-like, gently curved structure; external leaflet symmetrical,
striated, and with a slight apical point; 3 or 4 accessory setae present, rather
long, blade-like, with 1 or more retrorse apical processes; distimere normal
in shape, in Some specimens with a few very fine retrorse spines on the
convex surface (not illustrated); lateral plate of the phallosome with the internal
process rather broad, gently curved and pointed, not exceeding the apex of the
dorsal process; dorsal process with a spiculate apical knob and with 10 or
more teeth on the lateral margin.
LARVA. (Figure 29A,B). Head. Antenna with a narrow, dark basal
ring and darker beyond insertion of hair 1-A; head hair 4-C single (occasionally
bifid), simple; 5, 6-C bifid, pectinate, subequal in length (5-C may occasionally
be trifid); 16, 17-C represented by relatively well developed bifid spicules; a
band of very small tubercles curving around the anterior margin of the eye.
Thorax. Integument predominantly glabrous, but with rather distinctive
Spiculation on the anterolateral margins of the prothorax; 3-P single, pecti-
nate, shorter and slenderer than 1,2-P; 4-P usually bifid, pectinate (oc-
casionally trifid); 5,6-P single, pectinate; 7-P usually bifid, pectinate, but
sometimes with from 1 to 4 branches; 8-P single, pectinate; 14-P usually
bifid, simple, occasionally trifid. Abdomen. Integument glabrous; comb
consisting of from 30 to 50 scales arranged in a broad, somewhat triangular
patch; the posterior row of scales pointed and fringed with fine spines, the
other scales fan-shaped; siphon index variable, ranging from 7:1 to 11:1
(average, 8.3:1; in Malaya the range has been reported by Colless (1965) as
6. 7:1 to 8.8:1 with an average of 7.7:1); 3 pairs of subventral tufts present;
individual tufts usually 3 or 4 branched, but may range from 2 to 7 branches,
their length approximately twice the width of the siphon at the point of
insertion; pecten consisting of from 12 to 16 teeth restricted to the basal
third or less of the siphon; individual pecten tooth with a fine distal spine and
from 4 to 6 graded lateral barbs, the proximal barbs usually more prominent
and thicker than the distal barbs. )
TYPE DATA. Lectotype male (terminalia mounted and attached to
pin; antenna slide-mounted) of minor from Ulu Klang, Kuala Lumpur, Malaya
in the British Museum. The type locality of nolledoi is Kolambugan, Lanao,
Philippines, but the type specimen has been lost. The type locality of
bicornutus is Dawna Hills, near Kawkarech, Lower Burma, but the location
of the type specimen is unknown.
DISTRIBUTION. In THAILAND, this species has been studied
from: Chon Buri, Nakhon Ratchasima, Phattalung, Prachin Buvi, Ranong,
Songkhla, Tak, Tvat, and Trang. It is also known from MALAYA, BURMA,
and the PHILIPPINES.
The following specimens have been examined from Thailand during
the course of this study: 22 males, 12 with associated larval and pupal skins;
Bram: Genus Culex in Thailand 99
7 females, 4 with associated larval and pupal skins; 15 larvae.
TAXONOMIC DISCUSSION. The distinctive larval comb clearly
separates this species from other members of the mammilifer group; how-
ever, the adult male demonstrates affinity to pbeytoni. The relationship of
minor to minor form bicornutus will be discussed under the form.
BIOLOGY. Larvae of minor are most frequently collected in associa-
tion with bamboo habitats, but have also been collected in Thailand from tree
holes, rock pools, pandanus axils, stream pools, and root holes in jungle
areas. Collections were made during March, August, September, and
October. Nothing is known of the adult habits.
FORM BICORNUTUS
(Figure 29A')
FEMALE. Indistinguishable from minor (typical form).
MALE. Indistinguishable from minor (typical form) except for the
following character: proboscis generally slightly expanded distad of the
joint, with a distinct and characteristic ventrolateral series of rather long,
curved hairs on the expanded portion.
LARVA. Virtually indistinguishable from minor (typical form); the
individual pecten tooth has graded lateral barbs (figure 29A'), the basal ones
usually not distinctly more prominent and thicker than the distal ones.
DISTRIBUTION. In THAILAND, this form has been studied from
Chiang Mai, Chon Buri, Chumphon, Kanchanaburi, Lampang, Mae Hong Son,
Nakhon Nayok, Nakhon Ratchasima, Narathiwat, Phattalung, Pyvachin Buyvi,
Prachuap Khiri Khan, Ranong, Songkhla, Tak, Tvat, and Trang. It has also
been reported from MALAYA, western INDIA, lower BURMA, and Hungtow
Island, TAIWAN. Specimens have been examined from Con Son, SOUTH VIETNAM**.
The following specimens have been examined during this study: 75
females with associated larval and pupal skins, 87 males with associated
larval and pupal skins, 111 larvae, and 69 males and 46 females which
probably represent this form.
TAXONOMIC DISCUSSION. Edwards (1932) treated bicornutus as a
synonym of mammilifer; subsequently Colless (1965) properly removed
bicornutus from the erroneous synonymy and, although recognizing the close
anatomical and biological similarity with minor, elevated bicornutus to
species rank. Three primary characteristics (and several variable secondary
features) were presented by Colless to support his decision: (1) the presence
of a ventrolateral series of rather long, curved hairs on the expanded portion
of the male proboscis in bicornultus, as opposed to the absence of these hairs
in minor; (2) the pecten tooth of bicornutus with graded lateral barbs in
contrast to the condition in minor in which the proximal barbs are usually
more prominent and thicker than the distal barbs; and (3) the larval habitats
of bicornutus usually close to the ground and avoiding bamboo, those of minor
frequently in association with bamboo or tree holes.
Examination of specimens from Thailand strongly suggests at least
some area of overlap in these 3 main distinguishing characteristics. Although
the condition of the proboscis is clearly distinctive in most instances, 5 male
Specimens were examined which exhibited a somewhat intermediate condition
in which the ventrolateral hairs were present, but weakly developed despite
100 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
the presence of typically bDicornutus pecten teeth on the associated larvae.
The shape of the pecten teeth is quite variable, at least in specimens from
Thailand. It is not uncommon to see males with well developed ventrolateral
hairs on the proboscis but with both types of teeth in the pecten, or conversely,
to see males without the ventrolateral hairs but with a typically bicornutus
pecten. Of 74 collections of bicornutus containing individual rearings, 60
were made from habitats close to the ground; however, 3 collections were
made from bamboo internodes, 3 from pandanus axils, and 8 from tree holes:
of 16 collections of minor containing individual rearings, 9 were made from
habitats associated with tree holes or bamboo, and 7 were made from rock
pools, or on one occasion, a stream pool. It is therefore apparent that eco-
logical separation of the 2 forms in Thailand is not as distinctive as suggested
by Colless (1965) from data accumulated in Malaya.
Considering the above data from Thailand, it is felt that bicornutus
and minor should not be recognized as distinct species at the present time,
but that their close relationship could best be expressed by accepting bDicornutus
as a recognizable form of minor. Future ecological investigations and sibling
rearings would undoubtedly contribute much to a more refined and definitive
interpretation of this interesting complex.
BIOLOGY. Larvae of form bicornutus have been most frequently
found in habitats close to the ground. Included among these are: rock pools
and holes, stump holes, clay pots, coconut shells, tin containers, elephant
prints, stream pools, a spring, seepage pools, and sumps. As indicated
above, specimens have also been collected less frequently from bamboo
internodes, pandanus axils, and tree holes. Collections have been made
during every month of the year except December and February. Nothing is
known of the adult habits.
CULEX (LOPHOCERAOMYIA) PEYTONI BRAM AND RAT TANARITHIKUL 1967
(Figure 30)
Culex (Lophoceraomyia) peytoni Bram and Rattanarithikul 1967, Proc. ent.
Soc. Wash. 69: 7 (o*, 2, L*).
The adult male may be recognized by the presence of only a few long
hairs on flagellomere V, an internal tuft of at least 5 strongly sigmoid setae
on flagellomere VII, a tuft of strong, dark setae on flagellomere IX, and by
the internal rod of the subapical lobe of the basimere slightly expanded sub-
apically, followed by a rather broad, gently curved apex. The fourth stage
larva possesses a long siphon with 4 pairs of strong, long, subventral tufts;
the thoracic integument is covered with fine spicules, but the abdominal
integument is mostly smooth; the lateral denticles of the individual pecten
teeth are very broad proximally.
FEMALE. Thorax. Integument of the pleuron pale, without distinct
patches of scales; 1 lower mesSepimeral bristle present. Abdomen. Terga
dark brown; sterna somewhat lighter.
MALE. Head. Proboscis with a series of 6 or more short, baso-
ventral bristles. Antenna. (Figure 30D). Flagellomere V with a tuft of yel-
Figure 30. C. (Lophoceraomyia) peytoni. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male antennal
flagellomeres V through IX.
—_>
Genus Culex in Thailand
Bram:
“JL HWANG
C. (Lophoceraomyia) peytoni
102 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
lowish, unmodified hairs which reach to the tuft of IX; VI with a tuft of 4
strong, bent setae and 2 fine, straight setae; VII with a similar but shorter
tuft of 5 or more sigmoid setae; VII with an internal tuft of from 6 to 9
gently curved, dark brown setae; IX with a small internal tuft of 3 strong,
rather long, dark setae. Terminalia. (Figure 30C). Basimere with a
straight row of 4 or 5 strong, submarginal setae; subapical lobe of the basi-
mere with the internal rod slightly expanded subapically, then bent and with
a rather broad, lightly sclerotized apex; the central and external rods sub-
equal in length, sharply hooked; accessory processes narrow; internal leaf-
let rod-like, gently bent; external leaflet slender, striate; distimere with l
or 2 additional setae on the proximal third, and with minute annulations on
the convex surface of the distal third; lateral plate of the phallosome with the
internal process small, recurved, and pointed, not exceeding the apex of the
dorsal process; dorsal process with a subapical knob and 5 short denticles on
the lateral margin.
LARVA. (Figure 30A,B). Head. Antenna with a narrow, dark basal
ring and progressively darker beyond the insertion of hair 1-A; head hair 4-C
single, simple; 5,6-C bifid, pectinate; 16,17-C represented by minute spicules
which are easily overlooked even in good preparations. Thorax. Integument
covered with a sparse pattern of fine and rather long spicules; 3-P single,
pectinate, shorter and slenderer than 1,2-P; 4-P bifid, pectinate; 5,6-P
single, pectinate; 7-P single or bifid, pectinate; 8-P single, pectinate; 14-P
bifid, simple. Abdomen. Integument glabrous; hair 6-I trifid; comb consist-
ing of a broad, triangular patch of from 35 to 50 elongate, fan-shaped scales;
siphon index variable, ranging from 6.6:1 to 9.2:1 (average, 8:1); 4 pairs of
subventral tufts inserted in a line on the siphon; individual tufts with from 3 to
) branches, their length greater than the width of the siphon at the point of
insertion; pecten consisting of from 10 to 14 teeth restricted to the basal fourth
of the siphon; individual pecten tooth with an extended, fine distal spine, and
from 6 to 10 lateral barbs, the basal barb (sometimes the proximal 2 or 3
barbs) very large and rather rounded apically.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia and antenna slide-mounted) from Phattalung, Thailand in the U. S.
National Museum.
DISTRIBUTION. In THAILAND, this species is known from:
Chanthaburi, Chiang Mai, Chumphon, Narathiwat, Phattalung, Ranong, Sava
Buri, and Trang.
During this study 16 individual rearings have been examined as well
as 4 additional males and 70 larvae.
TAXONOMIC DISCUSSION. C. peytoni falls within the traubi-ganapathi
Species complex; its anatomical affinity is closest to ganapathi, but the larval
biology is rather intermediate. The adult male may be separated from
ganapathi primarily by the presence of a tuft of modified setae on flagello-
mere IX and by the presence of 1 or 2 setae on the proximal third of the disti-
mere of the terminalia; the male differs from fvaubi and spiculosus by the
presence of a tuft of long, narrow, tapering scales on flagellomere V. The
fourth stage larva may be distinguished from ganapathi by the form of the
pecten tooth in which the lateral denticles are very broad basally and by the
fact that head hair 1-C is never expanded on the basal half as frequently
found in ganapathi; the fourth stage larva is distinguished from tvaubi and
spiculosus by the glabrous abdominal integument and undeveloped median
caudal filament.
Bram: Genus Culex in Thailand 103
BIOLOGY. Larvae of peytoni have been collected in primary rain
forests from tree holes, bamboo internodes, and bamboo stump holes, and 1
collection was made from arock hole. Collections were made during the
months of January, May, and July through November. Habits of the adults
are unknown.
CULEX (LOPHOCERAOMYIA) PHOLETER BRAM AND RATTANARITHIKUL 1967
(Figure 31)
Culex (Lophoceraomyia) pholeter Bram and Rattanarithikul 1967, Proc. ent.
Soc. Wash. 69: 13 (o*, ?, L*).
The adult male is distinguished by flagellomere V possessing a
conspicuous tuft of from 1 to 4 long, broad scales, by the presence of 3 long,
prominent submarginal setae on the basimere, and by the shape of the internal
process of the lateral plate of the phallosome. The fourth stage larva exhibits
a long siphon with 4 pairs of long, rather fine, usually bifid subventral tufts
and thoracic hair 8-P branched, simple, and reduced in size.
FEMALE. Thorax. Integument of the pleuron pale, without distinct
patches of scales; 1 lower mesepimeral bristle present. Abdomen. Terga
dark brown; sterna very pale.
; MALE. Head, Proboscis with a series of 6 or more short, baso-
ventral bristles. Antenna. (Figure 31D). Flagellomere V with from 1 to 4
very broad scales which taper to a sharp point and extend to the tuft of VIII,
and several more narrow, but equally long setae; VI with a tuft of 6 or more
strong setae which are strongly recurved; VII with a tuft of 4 or more shorter,
strongly kinked setae; VIII with a tuft of 5 or more strong, broad, dark setae
which exhibit a prominent median bend; IX with an internal tuft of from 3 to 5
long, straight, strong setae. Terminalia. (Figure 31C). Basimere with 3
strong, Submarginal setae inserted in a straight row; subapical lobe of the
basimere with the internal rod tapering to a sharp point and shorter than the
central and external rods which are subequal in length and broadly bent at
the apex; internal leaflet rod-like, fine; external leaflet narrow, bluntly
rounded and striated; accessory processes narrow, setae-like; distimere
with minute annulations on the apical third of the convex surface; internal
process of the lateral plate of the phallosome small, gently curved, and
pointed, not exceeding the apex of the dorsal process; dorsal process with
an apical knob and approximately 7 short denticles on the lateral margin.
LARVA. (Figure 31A,B). Head, Antenna with a narrow, dark
basal ring and progressively darker beyond insertion of hair 1-A; head hair
4-C single, simple; 5, 6-C bifid, pectinate; 16,17-C represented by small,
but distinct spicules. Thovax. Integument glabrous; 3-P single, pectinate,
shorter and slenderer than 1,2-P; 4-P bifid, pectinate; 5,6-P single, pecti-
nate; 7-P bifid, pectinate; 8-P with 2 or 3 branches, simple, very short and
fine; 14-P bifid or trifid, simple. Abdomen. Integument glabrous; hair 6-I
bifid; comb consisting of a broad, triangular patch of from 60 to 80 elongate,
fan-shaped scales; siphon index variable, ranging from 6:1 to 7:1; 4 pairs of
subventral tufts inserted in a line on the siphon; individual tufts usually
bifid (occasionally trifid or single), their length greater than the width of the
Siphon at the point of insertion; pecten consisting of from 8 to 11 teeth
restricted to approximately the basal third to fourth of the siphon; individual
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
104
4
4
eC,
ay 7 We a
op N i “ :
ee eS =e
SSS 9
ZZ a 5
- X
C. (Lophoceraomyia) pholeter
Bram: Genus Culex in Thailand 105
pecten tooth with a prominent distal spine and from 7 to 10 lateral barbs, the
proximal 2 barbs very coarse, the distal barbs fine.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia and antenna slide-mounted) from Chon Buri, Thailand in the U. 8.
National Museum.
DISTRIBUTION. This species is known only from Chon Buri, THAI-
LAND. Three individual rearings were studied as well as 3 additional males
and 4 larvae.
TAXONOMIC DISCUSSION. Due to the presence of a tuft of conspicuous,
broad scales on flagellomere V, pholeter appears to have its closest affinity
to mammilifer; however, it may be separated from the latter species by having
only 3 prominent submarginal setae on the basimere and the internal process
of the lateral plate of the phallosome does not project beyond the apex of the
dorsal process. The fourth stage larva is also similar to mammulifer, but
can be recognized by the presence of 4 pairs of subventral tufts on the siphon;
the form of thoracic hair 8-P is very distinctive, and abdominal hair 6-I is
bifid rather than 3 or 4 branched.
BIOLOGY. Larvae of this species have been collected exclusively
from small crab holes in secondary rain forests. Collections were made
during the months of June, August, and October. The habits and biology of
the adults are unknown.
CULEX (LOPHOCERAOMYIA) SPICULOSUS BRAM AND RATTANARITHIKUL 1967
(Figure 32)
Culex (Lophoceraomyia) spiculosus Bram and Rattanarithikul 1967, Proc. ent.
Soc. Wash. 69: 3 (o*, 2, L*).
The adult male may be recognized by the absence of conspicuous tufts
on flagellomeres V and IX, by the submarginal setae of the basimere arranged
in a regular row, and by the internal process of the lateral plate of the phallo-
some not projecting beyond the dorsal process. The fourth stage larva is
characterized by possessing a long siphon with 4 pairs of strong, long, finely
pectinate, subventral tufts, having head hair 5-C bifid or trifid, 16,17-C
absent, and having the thoracic and abdominal integument covered with numerous,
prominent spicules.
FEMALE. Thorax. Integument of the pleuron pale, tinged with green
in Some specimens; 1 lower mesepimeral bristle present. Abdomen. Terga
dark brown; sterna slightly lighter.
MALE. Head. Proboscis with a series of 6 or more short, baso-
ventral bristles. Antenna. (Figure 32D). Flagellomere V with an incon-
spicuous group of from 6 to 8 very narrow, acute setae whose apices do not
Figure 31. C. (Lophoceraomyia) pholeter. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
sis of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
106 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
= Ze —7
=
3 ———
——S
a :
ew
etl
@ Vike
C.(Lophoceraomyia) spiculosus
Bram: Genus Culex in Thailand 107
reach beyond the tuft of VU; VI with a tuft of approximately 7 strong, some-
what twisted setae; VII with a similar tuft as well as 8 short, curved setae;
VIU with a tuft of long, rather straight, broad, dark setae; [X without a tuft
of obviously modified setae. Terminalia. (Figure 32C). Basimere with
from 6 to 8 strong submarginal setae inserted in a straight row; subapical
lobe of the basimere with the 3 rods subequal in length and width, gently
curved and pointed; internal leaflet rod-like, broad and bent; external leaflet
broad, symmetrical, pointed apically and striate; accessory processes nar-
row, setae-like; distimere normal, without apical annulations; lateral plate
of the phallosome with the internal process distinctly curved and pointed, not
projecting beyond the apex of the dorsal process; dorsal process with an
apical knob and approximately 10 short denticles on the lateral margin.
LARVA. (Figure 32A,B). Head. Antenna with a narrow, dark basal
ring and progressively darker beyond insertion of hair 1-A; head hair 4-C
single, simple; 5-C bifid or trifid, pectinate; 6-C bifid, pectinate; 16,17-C
absent. Thorax. Integument densely spiculose; 3-P single, pectinate,
shorter and slenderer than 1,2-P; 4-P bifid, pectinate; 5,6-P single, pecti-
nate; 7-P bifid, pectinate; 8-P single, pectinate; 14-P bifid, simple. Abdo-
men. Integument spiculose, but with the spicules not as prominent as those
of the thorax; hair 6-I trifid; comb consisting of from 35 to 45 fan-shaped
scales arranged ina broad, triangular patch; siphon index variable, ranging
from 7:1 to 11:1 (average, 8.8:1); median caudal filament short but distinct; 4
pairs of subventral tufts inserted in a line on the siphon; individual tufts with
from 2 to 5 branches which are finely pectinate, their length greater than the
width of the siphon at the point of insertion; pecten consisting of from 11 to
14 teeth restricted to approximately the basal fourth of the siphon; individual
pecten tooth with a prominent distal spine and from 6 to 8 lateral barbs, the
proximal 2 or 3 barbs very coarse; saddle with fine spicules on the distal
margin.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia and antenna slide-mounted) from Tak, Thailand in the U. S.
National Museum.
DISTRIBUTION. This species is known only from Chiang Mai, Nakhon
Nayok, and Tak, THAILAND. During this study 11 individual rearings have
been examined as well as 6 males, 3 females, and 50 larvae.
TAXONOMIC DISCUSSION. This species is a member of the traubi-
ganapathi species complex and demonstrates closest affinity, both in anatomi-
cal features and larval biology to fvaubi. It may be distinguished from traubi
by the internal process of the male phallosome which does not project beyond
the dorsal process; in the larva, hair 2-VIII of the abdomen is single in
spiculosus as opposed to bifid in fraubi. Additional distinguishing features
may be seen in the male antennae.
Figure 32. C. (Lophoceraomyia) spiculosus. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
thos
108 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
BIOLOGY. Larvae of spiculosus have been collected from large tree
holes in primary rain forests. There have also been 2 collections from a
bamboo stump and a bamboo internode. Collections were made during July,
August, and September. Nothing is known of the adult biology or habits.
CULEX (LOPHOCERAOMYIA) TRAUBI COLLESS 1965
(Figure 33)
Culex (Lophoceraomyia) traubi Colless 1965, J. med. Ent. 2: 295 (o*, 2, L*).
The adult male may be distinguished by the single row of submarginal
setae on the basimere, the absence of a tuft of modified setae on antennal
flagellomere IX, by the tuft of short, narrow, acute scales on flagellomere
V, and by the long internal process of the lateral plate of the phallosome.
The fourth stage larva exhibits 4 pairs of strong subventral tufts on the siphon,
the thoracic and abdominal integument is covered with numerous spicules,
and head hair 5-C is usually 3 or 4 branched.
FEMALE. Thorax. Integument of the pleuron dull brown, without
distinct patches of pale scales; 1 lower mesepimeral bristle present. Addo-
men. Terga dark brown; sterna somewhat paler.
MALE. Head. Proboscis without distinctive dorsal or ventrolateral
setae, but with a series of 6 or more short basoventral bristles; length of the
palpus often less than that of the proboscis. Antenna. (Figure 33D). Flagel-
lomere V with a tuft of golden, narrow, acute scales whose apices reach to
the tuft of VW; VI with a tuft of 10 or more strong, gently sigmoid scales
which have filamentous apices and several straight, narrow setae; VII witha
tuft of 7 or more narrow, short, sharply bent scales followed by 3 or 4
stronger, longer and kinked scales; VIII with an internal tuft of from 3 to 5
strong, darly pigmented, gently curved scales; [IX without modified scales or
setae. Terminalia. (Figure 33C). Basimere with 6 or 7 strong submarginal
setae usually inserted in a line, but often in an irregular row; subapical lobe
of the basimere with the internal rod very slightly expanded subapically, then
pointed, slightly shorter than the other rods; central and external rods sub-
equal in length, close to the internal rod and gently hooked; internal leaflet
rod-like, expanded and bent at the basal third; external leaflet rather broad,
symmetrical, finely striate; 3 or 4 fine, subequal accessory setae present;
distimere normal in shape, without a distal crest or annulations; lateral
plate of the phallosome with the internal process rather narrow, gently
curved, and projecting considerably beyond the apex of the dorsal process;
dorsal process with a spinose apical knob and with 10 or more strong lateral
teeth.
LARVA. (Figure 33A,B). Head. Antenna with a narrow dark basal
Figure 33. C. (Lophoceraomyia) traubi. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
—pP
109
Genus Culex in Thailand
e
e
Bram
C.(Lophoceraomyia) traubi
110 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
ring; head hair 1-C darkly pigmented, usually tapering gradually to a point
but occasionally slightly swollen on the basal half and with accessory lateral
Spicules; 4-C single, simple; 5-C 3 or 4 branched, pectinate; 6-C bifid or
trifid, pectinate, considerably longer than 5-C; 16,17-C represented by ex-
tremely minute spicules which are occasionally branched. Thorax. Integu-
ment minutely spiculate; 3-P single, pectinate, shorter and slenderer than
1,2-P; 4-P bifid, pectinate; 5,6-P single, pectinate; 7-P bifid, pectinate,
occasionally single; 8-P single, pectinate; 14-P bifid, simple, relatively
long. Abdomen. Integument minutely spiculate; hair 6-I trifid; comb consist-
ing of from 35 to 50 fan-shaped scales arranged in a broad, triangular patch;
siphon index variable, ranging from 7:1 to 12:1 (average, 9.2:1); the siphon
and segment X conspicuous by the dark pigmentation; 4 pairs of subventral
tufts inserted in a line on the siphon; individual tufts with from 2 to 7 strong
branches of decreasing length from base to apex, the basal pair usually with
only 2 or 3 branches, its length usually about 4 times the width of the siphon
at the point of insertion; pecten consisting of from 11 to 13 teeth restricted to
less than the basal third of the siphon; individual pecten tooth with a fine
distal spine, several very fine distal lateral barbs, and 2 strong proximal
lateral barbs; saddle with strong spines on the distal margin.
TYPE DATA. Holotype male (terminalia, antenna, and associated
larval and pupal skins slide-mounted) from Ulu Gombak, Selangor, Malaya in
the Australian National Insect Collection, Canberra.
DISTRIBUTION. C. tvaubi has been studied from 4 THAILAND** col-
lections from: Nakhon Nayok, Ranong, and Trang. The species has also been
recorded from Selangor and Perak, MALAYA.
During this study 2 males and 2 females with associated larval and
pupal skins (1 of each being a paratype) were examined in addition to 5 larvae.
TAXONOMIC DISCUSSION. As indicated under ganapathi, this species
is a member of the species complex which includes ganapathi, spiculosus,
and peytoni. C. traubi may be separated from the other members of the
complex primarily on the basis of the modified setae on the flagellomeres of
the male and the distinctive male terminalia; in the larval stage the abdominal
Spiculation and chaetotaxy of the head are diagnostic.
BIOLOGY. In Thailand larvae have been collected exclusively from
tree holes during the months of September and October. In Malaya collections
were also made from water in bamboo. Little is known of the adult biology,
although Colless (1965) has collected adults sweeping foliage and suggested
that they may attack man on occasion.
CULEX (LOPHOCERAOMYIA) WILFREDI COLLESS 1965
(Figure 34)
Culex (Lophoceraomyia) wilfredi Colless 1965, J. med. Ent. 2: 297 (o'*).
The adult male is immediately recognized by the modified torus of the
antenna and by the distinctive lateral plate of the phallosome. The fourth
Stage larva exhibits 4 pairs of fine, rather weak, subventral tufts on the
Siphon, thoracic hair 14-P single and simple, and abdominal hair 2-X trifid.
FEMALE. Thorax. Integument of the pleuron dull brown, without
distinct patches of pale scales, but with a line of small, dull scales on the
lower sternopleuron; 1 lower mesepimeral bristle present. Abdomen. Terga
Bram: Genus Culex in Thailand 111
dark brown throughout; sterna somewhat paler.
MALE. Head Proboscis without conspicuous dorsal or later oventral
setae, but with 8 or more short, basoventral bristles. Antenna. (Figure
34D). Torus without the mammiliform tubercle normally found in the
mammilifer group, but instead, slightly swollen on the inner surface and
with a circumferential, almost slit-like depression; flagellomere V with a
tuft of from 5 to 10 narrow yellow setae which extend beyond the tuft on IX;
VI with a tuft of 12 or more darkly pigmented, strongly sigmoid setae which
possess filamentous apices; VII with a similar tuft of broader setae; VIII
with a tuft of 10 or more darkly pigmented, gently curved and hooked setae;
IX with 5 or more strong, straight, yellow scales and a secondary tuft of
fine, dark, shorter setae. Terminalia. (Figure 34C). Basimere with from
4 to 6 strong submarginal setae inserted in a line at the base of which is a
tuft of from 3 to 8 shorter, straighter setae; subapical lobe of the basimere
with the basal rods subequal in length; the internal rod terminating in a point
which may at times appear to be filamentous; central and external rods gently
hooked; internal leaflet very narrow, rather straight, almost rod-like; ex-
ternal leaflet symmetrical, finely striated; 6 accessory setae present, but
viewed from the external aspect only 3 are usually visible (as illustrated);
distimere normal in shape, without distal annulations, but occasionally with
an extra basal seta; lateral plate of the phallosome with the internal process
very short, denticulate, about 1/3 or less the length of the dorsal process;
dorsal process very broad, with the apical half covered with numerous strong
teeth.
LARVA. (Figure 34A,B). Head. Antenna with a narrow, dark basal
ring and darker beyond insertion of hair 1-A; head hair 4-C single, simple;
9,6-C bifid, pectinate, subequal in length (5-C occasionally trifid, at least on
1 side); 14-C with 5 or 6 branches; 16,17-C represented by minute, bifid
spicules (occasionally 1 of the pair may be single on one side). Thorax. In-
tegument glabrous; 3-P single, pectinate, shorter and slenderer than 1, 2-P;
4-P bifid, pectinate; 5,6-P single, pectinate; 7-P bifid, pectinate (occasional-
ly trifid, as illustrated); 8-P single, pectinate; 14-P single, simple. Abddo-
men, Integument glabrous; hair 6-I trifid; 2-X trifid; comb consisting of
from 40 to 60 fan-shaped scales arranged in a broad, triangular patch; siphon
index variable, ranging from 8:1 to 10.3:1 (average, 8.7:1); 4 pairs of fine,
subventral tufts inserted in an irregular line on the siphon; individual tufts
usually bifid, sometimes trifid, their length less than the width of the siphon
at the point of insertion; pecten consisting of from 9 to 12 teeth restricted to
approximately the basal third of the siphon or less; individual pecten tooth
with a rather strong distal spine and from 5 to 7 graded lateral barbs.
TYPE DATA. Holotype male (terminalia and antenna slide- mounted)
from ''? Malaya" in the Australian National Insect Collection, Canberra.
DISTRIBUTION. In THAILAND**, wilfredi has been collected from:
Ayutthaya, Chiang Mai, Lampang, Nakhon Nayok, Nakhon Ratchasima, and
Tak. The species is also known from the type locality, probably in MALAYA.
During this study the following material was examined: 63 males,
4 with their associated larval and pupal skins; 5 females with associated
larval and pupal skins; and 27 larvae.
TAXONOMIC DISCUSSION. Colless (1965) provisionally placed this
Species within the mammilifer subgroup of the mammilifer group. Discovery
of the larval stage in Thailand confirms this decision. In the adult male,
wilfredi is quite distinctive from all other members of the group; the unusual
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
112
Fig.34
=
=——>
C. (Lophoceraomyia) wilfredi
Bram: Genus Culex in Thailand 113
antennal torus, although noticeably swollen on the inner margin, does not ex-
hibit the mammiliform protuberance common to the group, and the phallo-
some of the male terminalia is exceptional. The fourth stage larva demon-
strates an affinity to fuscosiphonis, but is recognized by the pale siphon and
features of the chaetotaxy.
BIOLOGY. Although specific data were not available, the type series
of this species was associated with a collection of material bred from pitcher
plants. In Thailand, larvae were collected from elephant prints, stream
pools, a ground hole, a pond, a sump, anda tree hole, and adults have been
collected resting on foliage. Collections were made during July, September,
and December through April. Nothing is known of the adult habits.
SUBGENUS THAIOMYIA BRAM 1966
Culex (Thaiomyia) Bram 1966, Proc. ent. Soc. Wash. 68: 73. Type species:
Culex dispectus Bram.
Characterization of this subgenus is based exclusively on the single
species, dispectus, described below. Thaiomyia demonstrates its closest
affinity to the subgenus Culiciomyia and the females of the 2 subgenera can-
not be separated. The male terminalia of Thaiomyia could also easily be
included with the Culiciomyia, but the third segment of the palpus is without
the distinctive lanceolate scales on the ventral surface which are so charac-
teristic of the latter subgenus. The fourth stage larva of Thaiomyia completely
lacks a pecten and the ventral brush consists of 10 individual tufts of setae,
the basal 1 located between the grid and the saddle. The subgenus Thaiomyia
is known only from Thailand.
CULEX (THAIOMYIA) DISPECTUS BRAM 1966
(Figures 35, 36, and 37)
Culex (Thaiomyia) dispectus Bram 1966, Proc. ent. Soc. Wash. 68: 75 (c*,
Pitty P*).
Salient features of this species are presented under the subgeneric
discussion above.
FEMALE. In general a moderately sized species with overall dark
brown appearance and without striking characteristics. Head. (Figure 35B).
Proboscis and palpus uniformly dark scaled; decumbent scales of the vertex
Sparse, narrow and light brown medially, gradually becoming lighter and
broader towards the orbital line; erect scales dark brown. Thorax. (Figure
tnt neering erent rence cane te i,
Figure 34. C. (Lophoceraomyia) wilfredi. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral aspect
ae of the terminal abdominal segments of the fourth stage larva;
C, dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia; D, lateral aspect of male
antennal flagellomeres V through IX.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
114
tus
x
ia) dispec
fomyt
4
C.(Tha
Bram: Genus Culex in Thailand 115
35B, E). Integument of the scutum uniformly light brown, with faintly darker
stripes in the dorsocentral area which are bordered by nude areas; the
scutum covered with a uniform pattern of bronze-brown scales; acrostichal
bristles absent, except for a very few small dark bristles at the extreme
anterior margin; anterior dorsocentral, posterior dorsocentral, supraalar,
and prescutellar bristles prominent and well developed; scutellum with the
- integument and scales similar to that of the scutum and with the bristles
normal for the genus; integument of the pleuron uniformly creamy white,
occasionally with indistinct, faintly darker patterns at the postspiracular
plate, and the lower sternopleuron; distinct scale patches or scattered scales
absent from the pleuron; upper sternopleuron with approximately 4 large and
1 small bristle; posterior sternopleuron with a series of from 5 to 10 bristles;
lower mesSepimeron with 1 strong and occasionally 1 weak bristle. Wing.
(Figure 35C). Legs. (Figure 35D). Anterior surface of the hind femur
dark, but with a variable line of light scales on the anteroventral margin;
hind tibia and tarsus as well as the fore and mid legs uniformly dark brown.
Abdomen. (Figure 35F). Terga completely dark brown; sterna covered with
a uniform pattern of light brown scales.
MALE. Similar in general appearance to the female except as noted
below. Head. (Figure 35A). Proboscis with an apparent median joint, at
which is inserted a ventral tuft of dark setae; length of the palpus exceeding
the proboscis by approximately the fourth and fifth segments; palpal segment
Iii without distinctive, lanceolate scales on the ventral surface; antenna
normal for the genus, slightly shorter than the length of the proboscis. Ad-
domen. Terga basically dark brown, but beginning with tergum IV, a basal
white spot is present which does not extend to the lateral margins and is
triangular in shape. Tervminalia. (Figure 36C). Subapical lobe of the
basimere with 3 strong rods and a strong seta on the proximal division and
2 strong setae on the distal division; distimere normal in shape, usually
with a crest of 1 or 2 barbs on the convex surface and with several short
setae on the basal third; lateral plate of the phallosome simple, witha
strong, but short basal tooth and approximately 8 lateral denticles; proctiger
well developed, crowned with a tuft of strong spines, and with 3 cercal setae
present; basal sternal process absent.
PUPA. (Figure 36A,B).
LARVA. (Figure 37). Head. Antennal shaft constricted beyond in-
sertion of hair 1-A; hairs 2-6-A inserted at the apex of the shaft; head hair
1-C filamentous, its length approximately 3/4 the distance between the bases
of the pair; 4-C simple, single or double; 5-C 12 branched, pectinate; 6-C
with approximately 9 branches, pectinate; 16,17-C represented by minute
spicules. Thorax. Integument glabrous; 1-P trifid, pectinate; 2-P single,
pectinate; 3-P with 2 or more branches, pectinate, considerably shorter and
Slenderer than the individual branches of 1, 2-P; 4-P trifid, pectinate; 5,6-P
Figure 35. C. (Thaiomyia) dispectus. Adult habitus: A, lateral aspect of
the male head; B, lateral aspect of the female head and thorax;
C, dorsal aspect of the female wing; D, anterior surface of
the female legs; E, dorsal aspect of the female scutum and
scutellum; F, dorsal aspect of the female abdomen.
the
116 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig. 36
ome eX
we
9
\
J.L. HWANG
C.(Thaiomyia) dispectus
Bram: Genus Culex in Thailand 117
single, pectinate; 7-P trifid, pectinate; 8-P very short, simple, dendritic;
14-P single, simple. Abdomen. Integument glabrous; comb consisting of
from 20 to 30 fan-shaped scales arranged in a broad, triangular patch;
siphon index variable, ranging from 3:1 to 4:1, rather broad basally and
tapering to a truncate apex; 4 pairs of subventral tufts inserted in a line,
their length greater than the width of the siphon at the point of insertion; in-
dividual siphon tufts with from 3 to 6 simple branches; pecten absent; saddle
completely ringing segment X; ventral brush consisting of 10 individual tufts
of setae, the basal tuft usually inserted between the grid and the saddle.
TYPE DATA. Holotype male with associated larval and pupal skins
and terminalia slide mounted from Tak, Thailand in the U. S. National
Museum.
DISTRIBUTION. This species is known exclusively from THAILAND
from: Chiang Mai, Ranong, and Tak. During this study the following material
was examined: 29 females and 27 males (23 of which were individual rearings
with associated larval and pupal skins), and 56 larvae.
TAXONOMIC DISCUSSION. As indicated in the subgeneric discussion
above, this species is similar to species of the subgenus Culiciomyia, but
features of the adult male and fourth stage larva clearly suggest a separate
erouping. It is the only species of the genus Culex known from Southeast
Asia which lacks a pecten in the fourth stage larva.
BIOLOGY. Larvae have been collected on 4 occasions from bamboo
internodes and stumps in a primary rain forest environment. Collections
were made during the months of August and September, and associated species
taken with dispectus included Armigeres flavus, A. subalbatus, Orthopodomyia
sp., Uranotaenia sp., and C. (Lophoceraomyia) sp. Nothing is known of the
adult habits or biology.
SUBGENUS CULICIOMYIA THEOBALD 1907
Trichorhynchus Theobald 1905, J. Bombay nat. Hist. Soc. 16: 241 (preoc-
cupied by Balbiani 1887). Type species: Trichorhynchus fuscus
Theobald.
Culiciomyia Theobald 1907, Mon. Cul. 4: 227. Type species: Culiciomyia
inornata Theobald (Edwards 1912, Bull. ent. Res. 3: 33).
Neomelanoconion Theobald 1907, Mon. Cul. 4: 514. Type species: Culex
vima Theobald.
Pectinopalpus Theobald 1910, Ann. Mag. nat. Hist. 5: 375. Type species:
Pectinopalpus fuscus Theobald.
Trichorhynchomyia Brunetti 1912, Rec. Indian Mus. 4: 477. wee Species:
Trichorhynchus fuscus Theobald.
The adult female may be recognized by the presence of 1 or 2 lower
mesepimeral bristles, the unbanded proboscis, the absence of scale patches
on the pleuron, the acrostichal bristles not developed, and the scaling of the
Figure 36. C. (Thaiomyia) dispectus. A,B, dorsoventral aspects of the
pupa; C, dorsal aspect of the male terminalia.
118 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.37
10
y 12
V4
MP
9
10 32
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a Wty,
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att \
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167 Se
ye 9
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S
SS
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7
C. (Thaiomyia) dispectus
Bram: Genus Culex in Thailand 119
scutum very dense and smooth in appearance. The adult male may be
recognized by the above characters as well as the presence of a linear series
of lanceolate scales on the ventrolateral surface of the third segment of the
palpus, and the characteristic terminalia. The fourth stage larva is distin-
guished by the pecten prominent and restricted to the basal half of the siphon,
and the ventral brush consisting of 8 hair tufts inserted within the grid.
| FEMALE. Usually small to medium sized species. Head. Proboscis
unbanded; vertex with broad, decumbent scales at the orbital line, narrowing
towards the occiput. Thorax. Scaling pattern of the scutum very dense,
smooth in appearance; acrostichal bristles not developed except at extreme
anterior end and rarely weakly near prescutellar space; pleuron without
distinct scale patches; usually 1 or 2 strong lower mesepimeral bristles
present; wings with the dorsal plume scales short and rather broad; legs
without pale bands on the femur, tibia or tarsus. Abdomen. Terga with or
without basal or apical banding patterns.
MALE. Proboscis with a median tuft of several long, ventral hairs;
palpus longer than the proboscis, with a single row of flattened, distinctively
lanceolate scales on the ventral surface of segment III; antenna plumose,
slightly shorter than the proboscis. Terminalia. Subapical lobe of the basi-
mere with strongly developed setae and with or without a leaf-like seta; some
Species have a prominent sternal accessory apical lobe bearing very long,
hairlike setae; distimere variable in shape; phallosome simple, the lateral
plate with a strong basal tooth and usually with a series of short denticles on
the lateral margin; proctiger with a dense tuft of fine dorsomesal spines and
prominent, spatulate lateroventral spines; cercal setae short, variable in
number; basal sternal process variable.
LARVA. Head. Hair 1-C either fine and filamentous or robust and
tapering gradually; 16,17-C present, represented by minute spicules. Thorax.
Integument glabrous or spiculose; hair 1-P single or bifid; 2-P always single;
3-P single or branched, the branches always much thinner than, and usually
about half the length of 1-P; 4-P single or branched. Abdomen. Integument
glabrous or spiculose; comb consisting of a triangular patch of from 25 to 70
small, fan-shaped scales; siphon variable in length and shape, with from 3
to 6 pairs of subventral tufts inserted in a line beyond the pecten; ventral
brush consisting of 8 hair tufts inserted on the grid; anal gills always longer
than the saddle.
PUPA. Without striking subgeneric characteriotins:
DISTRIBUTION. The subgenus Culiciomyia is represented principally
in the Old World tropics, although a few species extend to Japan and 3 species
are found in the South Pacific. Collections to date indicate that members of
the subgenus are distributed throughout THAILAND.
TAXONOMIC DISCUSSION. Diagnostic subgeneric and species charac-
ters of Culiciomyia in Thailand are indistinct in the adult female. On the
basis of female characters the subgenus appears most closely related to
Thaiomyia and Lophoceraomyia and can be recognized by dense, smooth
scaling of the scutum, whereas these scales in Lophoceraomyia are generally
Figure 37. C. (Thaiomyia) dispectus. Fourth stage larva: dorsoventral
view of the head, thorax and abdomen, and lateral aspect of
€ the terminal abdominal segments.
120 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
sparse and rough in appearance. Within the subgenus, females of several
species are extremely difficult to separate and conclusive determinations
must still depend upon characters of the male terminalia and fourth stage
larvae. The male terminalia of the subgenus are distinctive and are charac-
terized by the simple phallosome and features of the basimere and distimere.
The fourth stage larva can be separated from other subgenera by the ventral
brush and by the thoracic chaetotaxy. At the present time the fourth stage
larva exhibits the most reliable features upon which species determinations
may be based.
BIOLOGY. The immature stages of species of Culiciomyia in Thai-
land are found in a variety of habitats ranging from tree holes to running
creeks and swamps in tropical rain forests; however, the larvae are found
most frequently in temporary accumulations of water, usually with high
organic content. Host preferences and bionomics of the adult females are
virtually unknown, but species of this subgenus have rarely been collected
feeding on humans, even in areas where the mosquitoes are quite numerous.
Two species, pallidothorax and spathifurca, have been experimentally fed on
humans infected with Wuchereria bancrofti with subsequent isolation of
infective larvae from the mosquito. It is felt, however, that these species
play little if any part in the transmission of the disease in nature.
KEY TO SPECIES OF THE SUBGENUS CULICIOMYIA IN THAILAND -
ADULT FEMALES!
1, At least some of the abdominal terga with apical or basal pale
apts ee ae ee, Ome ee
2(1). Dorsal wing scales creamy white; apical light bands covering the
greater portion of each tergum; scutum covered with pale
BODIE a I eon ee a Ie oa termi (p. 157)
Dorsal wing scales ranging in color from reddish-brown to black;
basal pale bands covering not more than half of each tergum;
Brie With Gark Scales. Ga ee Oe Oe a ea
3(2). Integument of the pleuron uniformly pale brown or with, at most, a
faint light brown stripe from the anterior pronotal lobe to the
lippermiesepimerons. 166 es ea BOG bailyi (p. 123)
Integument of the pleuron with distinct dark markings...........
é.
4(3). Integument of the pleuron with a prominent, isolated, very dark brown
to black spot present on the upper meSepimeron; a light brown
spot is usually present on the upper sternopleuron...........
nigvopunctatus (p. 133)
Integument of the pleuron with a brown pattern stretching from the
prespiracular areas across the prealar area and terminating at
the upper mesepimeron; a light brown pattern also present on
1
The adult female of spiculothorax is unknown; the placement of barrinus is
presumptive.
e@
Bram: Genus Culex in Thailand 121
the upper sternopleuron.............. pallidothorax (p. 137)
thurmanorum (p. 159)
barrinus (p. 125)
5(1). Segment IV of the palpus over 3 times as long as II; in general,
a laree, “dark species: 86605 CU papuensis (p. 143)
Segment IV of the palpus about twice as long as III; in general, a
biiall, shter Species ee Var er ee fragilis (p. 129)
spathifurca (p. 151)
scanloni (p. 14'7)
KEY TO SPECIES OF THE SUBGENUS CULICIOMYIA IN THAILAND -
ADULT MALES
1. Distimere divided proximally into 2 unequal areas (the external arm
flattened, expanded, and denticulate on the distal half; the
internal arm slender, pointed, and sigmoid). . spathifurca (p. 151)
Distimere represented by a single, undivided arm............. 2
2(1). Distimere with an apical crest of at least 3 recurved spines on the
CONVEX SULINOG Coo TT ey ee ee 3
Distimere without a crest of spines on the convex surface........ 7
3(2). Abdominal terga with distinct, pale basal bands............... 4
ADdONITNAl TEP Ca UNDANGee OO ee ae Oe es Be ka 6
4(3). Subapical lobe of the basimere with the leaf-like seta very broad;
integument of the pleuron with a prominent, isolated, very dark
brown to black spot present on the upper meSepimeron.........
nigropunctatus (p. 133)
Subapical lobe of the basimere with the leaf-like seta very slender or
absent; integument of the pleuron without an isolated, dark brown
Spot on the upper mesepinieron 7... ee ee a ee a 5)
5(4). Basal rod on the subapical lobe of the basimere shorter and slenderer
than the other 2 rods, and not inserted laterally out of line;
slender pee like seta rather long and usually conspicuous.......
pallidothorax (p. 137)
Basal rod on the subapical lobe of the basimere as long as and
broader than the other 2 rods and inserted laterally out of line;
slender leaf-like seta short and usually inconspicuous.........
thurmanorum (p. 159)
barrinus (p. 125)
6(3). Subapical lobe very near the apex of the basimere; a prominent
sternal apical area present bearing a patch of very fine
Cee Pe ee papuensis (p. 148)
Ithe adult male of spiculothorax is unknown; the placement of barrinus is
presumptive.
122 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Subapical lobe distinctly below the apex of the basimere; no sternal
apical area with fine setae present.......... fragilis (p. 129)
scanloni (p. 147)
7(2). Phallosome with from 4 to 6 prominent, heavily sclerotized denti-
cles; all dorsal wing scales uniformly dark; abdominal terga with
BAGG) BONE DONE boii) asic eck ea de ae eaanete be id bailyi (p. 123)
Phallosome with at most faint indications of from 7 to 13 small,
lightly sclerotized denticles in addition to the large basal tooth;
all dorsal wing scales pale; abdominal terga with broad apical
Te Ue a teu go eosia es Week Wie termi (p. 157)
KEY TO SPECIES OF THE SUBGENUS CULICIOMYIA IN THAILAND -
FOURTH STAGE LARVAE
i. Siphon index approximately 30:1 or more; thoracic hair 4-P with
Pe a aiips ieee oso olan 4 wiehe le termi (p. ae
Siphon index 15:1 or less; thoracic hair 4-P bifid or PE assis to «
AUER Pe De INCL i is ied s csraielinis wen ayet edt 0 «4
Thoracic hair 1-P bifid, 2-P single, and 3-P either single or with
OO RN aii ele ec eeles Aes hae me Seb nin lee Um nciaeen ome WE lol w
3(2). Siphon with a false joint beyond the middle due to lack of sclerotiza-
fiom in an Irewular FING. 6 666i Bn eceia nigropunctatus (p. oe
Siphon without a false joint beyond the middle...............2.
4(3). Basal barb of each individual pecten tooth very robust, considerably
larger than the other lateral barbs; siphon index approximately
BE OR POR AR oe Wee or See Ses scanloni (p. 14'7)
Basal barb of each individual pecten tooth smaller than, or equal
to the other lateral barbs; siphon index approximately 6:1 or
Re PA a a uml el le vive
5(4). Head hairs 5,6-C with 3 or 4 branches; individual siphon tufts
Pe ink i hdd euek Whee aa dace: Wenmabeniel ni vateuerce ah spathifurca (p. 151)
Head hairs 5,6-C with from 5 to 7 branches; individual siphon tufts
Wi Oe I eee Oe OL ES fragilis (p. 129)
6(2). Thorax and abdomen with a series of stellate setae; abdominal
integument covered with a uniform pattern of round, platelet-
DRE SPICWOR eS ee ae EN thurmanorum (p. 159)
Thorax and abdomen without stellate setae; abdominal integument
without platelet-like Spicules 2.6. ee ee ee wae 8
7(6). Head hair 1-C robust, lightly pigmented, and frequently possessing
accessory lateral spicules.............. papuensis (p. 143)
head hair t-C tine and filamentous 46355 0 6 ee es 8
8(7). Thoracic integument with a conspicuous pattern of coarse spicules. . .
spiculothorax (p. 155)
Bram: Genus Culex in Thailand 123
Thoracic integument glabrous, granulose, or with an inconspicuous
patiern:.of minute spicules) .3.0 woe whe es ib ee Wiad ails ws 9
9(8). Siphon narrow throughout its length and gently curved distally; more
than 10 teeth present in the pecten; subventral tufts of the siphon
either sineioor bifid 6s ea bailyi (p. 123)
Siphon enlarged medially; less than 10 teeth present in the pecten;
subventral tufts of the siphon usually with 3 or 4 branches. .... 10
10(9). Siphon with a broad, dark, subapical band....... barrinus (p. 125)
piphon wepanded oe. ety eee Mee heuk Yad pallidothorax (p. 137)
CULEX (CULICIOMYIA) BAILYI BARRAUD 1934
(Figures 38, 39, and 40)
Culex (Culiciomyia) bailyi Barraud 1934, Fauna Brit. India, Diptera 5:
382 (o*, 2); Carter and Wijesundara 1948, Ceylon J. Sci. 23: 147
(L*),
The adult female is identified by the pale basal bands of the abdominal
terga and by the absence of distinct integumental markings. The adult male
is recognized by the absence of a crest of spines on the convex surface of the
distimere and the robust lateral plate of the phallosome. The fourth stage
larva is distinguished by the characteristic shape and length of the siphon,
the glabrous thoracic integument, and the chaetotaxy of the head and pro-
thorax.
FEMALE. In general, a small to moderately sized species with
overall light brown appearance and without striking characteristics other than
the basal banding of the abdominal terga. Head. (Figure 38E). Proboscis
and palpus uniformly dark scaled; decumbent scales of the vertex light brown
on the occiput, gradually becoming lighter towards the orbital line. Thorax.
(Figure 38B,E). Integument of pleuron uniformly pale or with a greenish cast
in Some specimens; 1 strong lower mesepimeral bristle present. Wing.
(Figure 38C). Legs. (Figure 38D). Anterior surface of hind femur dark, but
with a line of lighter scales on the ventral margin; hind tibia and tarsus as
well as the fore and mid legs completely dark. Abdomen. (Figure 38F).
Terga dark with rather broad, pale basal bands; sterna pale.
MALE. Head. (Figure 38A). Proboscis with an apparent median
joint, at which is inserted a ventral tuft of dark setae. Terminalia. (Figure
39C). Subapical lobe of the basimere somewhat flattened but well developed
near the apex of the basimere, with strong setae and rods, but without a
leaf-like seta; distimere normal, without a crest of spines on the convex
surface but undulating distally and with a hooked structure above the claw;
lateral plate of the phallosome with a very strong and heavily sclerotized
basal tooth and from 4 to 10 smaller lateral denticles.
PUPA. (Figure 39A,B).
LARVA. (Figure 40). Head, Antenna with a narrow, dark basal ring;
head hair 1-C filamentous, its length greater than half the distance between
the bases of the pair; 4-C single, bifid, or trifid, very finely pectinate; 5,6-C
with 3 or 4 branches, pectinate. Thorax. Integument superficially glabrous,
124 Contrib.
Amer. Ent.
Inst., vol. 2, no. 1, 1967
C. (Culiciomyia) bailyi
Bram: Genus Culex in Thailand 125
but critical examination reveals the presence of extremely minute spicules
(not illustrated); thoracic hair 1-P bifid, pectinate; 2-P single, pectinate;
3-P bifid, pectinate, shorter than 1,2-P; 4,7, 8-P bifid, pectinate; 5,6-P
single, pectinate; 14-P single, simple. Abdomen. Integument with minute
spicules similar to those of the thorax; comb consisting of approximately 35
fan-shaped scales arranged in a triangular patch of 3 irregular rows; siphon
index variable, ranging from 6.5:1 to 7.5:1, the siphon narrow throughout its
length and gently curved distally; 3 pairs of subventral tufts inserted in a
line beyond the pecten; individual tufts single or bifid, their length, at most,
only slightly greater than the width of the siphon at the point of insertion;
pecten consisting of from 15 to 18 teeth restricted to the basal third to fourth
of the siphon; individual pecten tooth with a strong apical point and 3 or 4
strong lateral barbs.
| TYPE DATA. Lectotype hereby designated: syntype male (terminalia
slide-mounted), 'TBM 1935-622, Virajpet, Coorg, S. India, VI1927, J. D.
Baily", in the British Museum.
DISTRIBUTION. In THAILAND**, bailyi has been studied from:
Chiang Mai, Lampang, Mae Hong Son, Nakhon Nayok, Nakhon Ratchasima,
Narathiwat, and Ranong. Previously it had been recorded only from the
type locality in INDIA and from Hakgala, CEYLON. During this study 26 in-
dividual rearings were examined as well as 31 females, 27 males, and 75
larvae.
TAXONOMIC DISCUSSION. This species demonstrates its closest
affinity to pallidothorax, and is related to a lesser degree to nmigropunctatus.
However, the adult female of bailyi may be distinguished from the other 2
species by the absence of distinct dark markings on the pleuron; the adult
male lacks a distinct crest of spines on the distimere of the male terminalia;
and the fourth stage larva exhibits thoracic hairs 1, 3-P branched as in
_ pallidothorax, but lacks the expanded siphon of the latter species.
BIOLOGY. Carter and Wijesundara (1948) reported collecting larvae
in January from a cement cistern at a height of 5, 600 feet in Ceylon. In
Thailand, larvae have been collected from sumps, open puddles, rock pools,
elephant hoof prints, and tree holes during the months of January through
May, August, September, and November. Habits and biology of the adults
are unknown.
CULEX (CULICIOMYIA) BARRINUS, N. SP.
(Figure 41)
The fourth stage larva is characterized by possessing a broad, dark,
subapical band on the siphon.
FEMALE, MALE (Figure 41C). Adults presumed to be this species
cannot be distinguished from thurmanorum.
Figure 38. C. (Culiciomyia) bailyi. Adult habitus; A, lateral aspect of
| the male head; B, lateral aspect of the female head and
<<— thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female
abdomen.
126 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.39
C.(Culiciomyia) baily/
Bram: Genus Culex in Thailand 127
LARVA. (Figure 41A,B). Head. Antenna without the usual darker
basal ring; hair 1-C filamentous, but rather broad basally, its length slightly
greater than half the distance between the bases of the pair; 4-C single,
simple; 5,6-C bifid, pectinate. Thorax. Integument superficially glabrous,
but when examined critically, with extremely minute, fine, pointed spicules
(not illustrated); hair 1-P bifid, pectinate; 2-P single, pectinate; 3-P with 3
or 4 branches, pectinate; 4, 7-P bifid, pectinate; 5,6-P single, pectinate; 8-P
bifid or trifid, pectinate; 14-P single, simple. Abdomen. Integument
glabrous; comb consisting of approximately 35 elongate, fan-shaped scales
arranged in a triangular patch; siphon index ranging from 5:1 to 6:1, the
Siphon expanded medially and with a conspicuous, dark, rather broad sub-
apical band; 4 or 5 pairs of subventral tufts inserted in a line beyond the
pecten; individual tufts with 3 or 4 branches, their length less than the width
of the siphon at the point of insertion; pecten consisting of from 4 to 6 teeth
restricted to the basal third of the siphon; individual pecten tooth with a
strong distal spine and from 5 to 9 fine lateral barbs.
TYPE DATA. Holotype: fourth stage larva from Ban Pa Chi, Mae
Hong Son, Thailand, 10 September 1966, S. Maniwongse, from elephant
hoofprints in a secondary rain forest, at 600 m. elevation, deposited in the
U. S. National Museum, No. 69512. Paratypes: 2 larvae with the same
data as the holotype, deposited in the U. S. National Museum, No. 69512.
DISTRIBUTION. This species is known only from THAILAND from
the type locality of Mae Hong Son and from Lampang. In addition to the type
Series, 10 larvae have been examined; 11 females and 7 males which are
presumed to be this species were reared from pupae collected in association
with the larvae, however, other closely related species of Culiciomyia were
also collected from the same habitats.
TAXONOMIC DISCUSSION. The presumptive adults of barrinus can-
not be separated from thurmanorum and are also extremely similar to
_ pallidothorax. The fourth stage larva is also extremely similar to
_ pallidothorax, but is immediately recognized by the broad, subapical dark
band on the siphon. Larvae of both barrinus and pallidothorax have been
collected from the same habitat.
BIOLOGY. Larvae of barrinus were first collected in March 1953
from puddles and elephant tracks at kilometer 114 on the Lampang to Phayao
road. Larvae were apparently not collected again until September 1966,
when 4 separate collections were made in Mae Hong Son. Each of the 1966
collections were made from elephant hoof prints. The collections from Ban
Mae Ho Nua and Doi Chang included larvae of pallidothorax, bailyi, thurmano-
vum, and Anopheles balabacensis. In each instance collections were made
at an elevation of 600 m. or above in a secondary rain forest. Nothing is
known of the habits of the adults.
Figure 39. C. (Culiciomyia) bailyi. A,B, dorsoventral aspects of the
< pupa; C, dorsal aspect of the male terminalia.
128 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
3
\
‘
C.(Culiciomyia) bailyi
/
Bram: Genus Culex in Thailand 129
CULEX (CULICIOMYIA) FRAGILIS LUDLOW 1903
(Figures 42, 43, and 44)
Culex fragilis Ludlow 1903, J. N. Y. ent. Soc. 11: 142 (¢, 9).
Trichorhynchus fuscus Theobald 1905, J. Bombay nat. Hist. Soc. 16: 242
(2*); Edwards 1922, Indian J. med. Res. 10: 472 (synonymy).
Culiciomyia inornata Theobald 1907, Mon. Cul. 4: 227 (o«*, 9*); Edwards
1922, Indian J. med. Res. 10: 472 (synonymy).
Culiciomyia ceylonica Theobald 1907, Mon. Cul. 4: 236 (<,?*); Edwards
1922, Indian J. med. Res. 10: 472 (Synonymy).
Culex graminis Leicester 1908, Cul. Malaya: 158 (“, 2); Edwards 1932, in
Wytsman, Genera Insect. fasc. 194: 199 (synonymy).
Culex (Culiciomyia) fragilis Ludlow: Barraud 1934, Fauna Brit. India,
Diptera 5: 385 (o, 2); Bohart 1945, Navmed 580: 76 (‘*); King and
Hoogstraal 1946, Proc. biol. Soc. Wash. 59: 148 (o'*, L*); Penn
1947, Pacif. Sci. 3: 72 (P*); Bick 1951, Pacif. Sci. 5: 417
(ecology); Bonne-Wepster 1954, Roy. trop. Inst. Amst., Spec. Pub.
111: 113 (o, 9, L*); Belkin 1962, Mosq. S. Pacif. I: 231 (o*, 2, P*,
L*); Delfinado 1966, Mem. Amer. ent. Inst. 7: 117 (o*, 9, L*, P*).
Due to the absence of distinctive anatomical features or color charac-
ters it is not presently possible to positively differentiate the adult female
from several other species in the subgenus. The adult male is recognized
on the basis of the terminalia in which the number and shape of setae on the
subapical lobe of the basimere are distinctive and by the absence of a promi-
nent sternal accessory apical lobe on the basimere. The fourth stage larva
may be distinguished by head hairs 5, 6-C with from 5 to 8 branches, by the
subventral tufts of the siphon bifid or trifid, by the shape of the pecten teeth,
and usually by the siphon index. _
FEMALE. A relatively small species without distinctive anatomical
features or color patterns. Head. (Figure 42B). Proboscis and palpus dark
scaled; palpal segment IV twice as long as III; decumbent scales of the vertex
light brown at the occiput and becoming lighter as they progress to the orbital
line; erect scales predominantly light brown, but several may appear creamy
white. Thorax. (Figure 42B,E). Integument of the pleuron light brown; 1
strong, lower mesepimeral bristle present. Wing. (Figure 42C). Legs.
(Figure 42D). Anterior surface of the hind femur dark with a pale stripe on
the ventral margin; hind tibia and tarsus uniformly dark, occasionally with
lighter scales randomly scattered, but without distinct patterns; fore and mid
legs with markings similar to those of the hind legs. Abdomen. (Figure 42F).
Terga dark to dusky brown, without pale bands or patterns; sterna somewhat
lighter.
MALE. Head. (Figure 42A). Proboscis with a median tuft of 2 or 3
fine setae on the ventral surface. Terminalia. (Figure 43E). Subapical lobe
of the basimere well developed, with 3 basal rods, 2 narrow, striated leaflets,
Figure 40. C. (Culiciomyia) bailyi. Fourth stage larva: dorsoventral
view of the head, thorax and abdomen, and lateral aspect of
<< the terminal abdominal segments.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
130
wT ;
ae
ve
oN
J. L. HWANG
C. (Culiciomyia ) barrinus
Bram: Genus Culex in Thailand 131
a broad, symmetrical, striated leaf, and a strong seta; distimere with a crest
of 10 or more recurved spines on the convex surface and a moderately promi-
nent hook opposite the claw; lateral plate of the phallosome with a strong,
heavily sclerotized basal tooth and from 6 to 10 evenly spaced, strong
denticles. :
PUPA. (Figure 43A, B).
LARVA. (Figure 44). Head. Antenna with a narrow, dark basal
band; head hair 1-C filamentous, its length slightly greater than half the
distance between the bases of the pair; 4-C single, simple; 5, 6-C with from
5 to 8 branches, pectinate. Thorax. Integument glabrous; hairs 1,2,3-P
Single, pectinate, with 3-P much shorter and slenderer than 1, 2-P; 4,7, 8-P
bifid, pectinate; 5,6-P single, pectinate; 14-P bifid, simple. Abdomen. In-
tegument glabrous; comb consisting of a triangular patch of from 25 to 40
fan-shaped scales; siphon index variable, averaging between 5:1 and 6:1; 3
subventral tufts inserted in a line on the siphon beyond the pecten; individual
tufts with 2 or 3 simple branches, their length greater than the width of the
siphon at the point of insertion; pecten consisting of from 10 to 13 teeth
restricted to the basal fourth to third of the siphon; individual pecten tooth
with a strong apical spine and about 4 strong, lateral barbs.
TYPE DATA. Lectotype male of fragilis (terminalia slide- mounted)
from Oras, Samar, Philippines in the U. S. National Museum. Holotype
female of fuscus from Peradeniya, Ceylon in the British Museum. Lectotype
of znornata hereby designated: syntype female, "B.M. 1907-29, Kuching,
Sarawak, Dr. Barker", in the British Museum. Lectotype of ceylonica here-
by designated: syntype female, 'B.M. 1907-29, Peradeniya, Ceylon,
2-1902"', in the British Museum. The type locality of graminis is Kuala
Lumpur, Selangor, Malaya, but the type specimen is non-existent.
Considering the lack of distinguishing features in the adult female and
the present inability to separate fragilis from scanloni in the adult female, it
is most unfortunate that it has been necessary to select female syntypes as
the lectotypes of zzornata and ceylonica. In both instances, however, the
syntype male lacked the terminalia and the rest of the specimen was not as
well preserved as in the syntype female.
DISTRIBUTION. This species was first reported from THAILAND by
Causey (1937) from "Haad Raj Samran".. Collections made during the course
of this study have been from: Nakhon Ratchasima, Prachuap kKhiri Khan,
and Ranong. This species has also been recorded from: PHILIPPINES,
INDIA, CEYLON, MALAYA, and INDONESIA. The records from New Guinea,
the Solomon Islands, and the Admiralty Islands (Belkin 1965), probably refer
to one or more distinct, but closely related species.
The following specimens have been examined from Thailand: 5 in-
dividual rearings, 13 females, 15 males, and 22 larvae.
TAXONOMIC DISCUSSION. C. fragilis demonstrates its closest af-
finity to scanloni but may be recognized by the differences in the fourth stage
larvae outlined in the key. The description of fragilis from the Solomon Islands
Figure 41. C. (Culiciomyia) barrinus. A, dorsoventral aspect of the head
and prothorax of the fourth stage larva; B, lateral aspect of the
a a terminal abdominal segments of the fourth stage larva; C, dorsal
aspect of the male terminalia.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
132
ja) fragilis
°
iciomy
°
(Cul
Bram: Genus Culex in Thailand 135
(1962) and specimens determined as this species by him in the U. S. National
Museum collection differ to a considerable degree from the species as it
occurs in Southeast Asia. In the male terminalia the structures of the sub-
apical lobe of the basimere and the denticles on the lateral plate of the phallo-
some show recognizable differences; in the fourth stage larva head hairs
5,6,16,17-C, the siphon index, the individual siphon tufts, and the shape of
the pecten tooth all differ consistently. It is therefore concluded (as Belkin
himself suspected), that the Solomon Islands form is not conspecific with
fragilis as here presented.
BIOLOGY. In Thailand, larvae of fragilis have been collected from
elephant tracks, ditches, and arock pool. Bick (1951) collected larvae of
fragilis in New Guinea primarily from artificial containers, followed by
frequence by coconut shells, puddles, pot holes, tree holes, ponds, spathes
and leaves, rock holes, ditches, and 1 collection from a running creek.
Water pH ranged from 5.0 to 7.0 and averaged 5.8. Sixty-two percent of the
collection sites were shaded, 64 percent of the sites had stagnant or polluted
water, and algae occurred in 63 percent of the habitats. King and Hoogstraal
(1946), also reporting from New Guinea, found larvae to be most commonly
- taken from temporary and semipermanent ground pools, especially with
stagnant, more or less foul or algae-filled water. Biology and habits of the
adults are unknown, but the females have never been reported attacking man.
CULEX (CULICIOMYIA) NIGROPUNCTATUS EDWARDS 1926
(Figures 45, 46, and 47)
Culiciomyia annulata Theobald 1907, Mon. Cul. 4: 231 (preoccupied in
Culex by Schrank 1776).
Culex nigropunctatus Edwards 1926, Bull. ent. Res. 17: 121 (new name for
annulata Theobald 1907, not Schrank 1776).
Culex (Culiciomyia) nigropunctatus Edwards: Barraud 1934, Fauna Brit.
India, Diptera 5: 383 (o¢*, 2, L); Bohart 1945, Navmed 580: 77 (<*);
Bohart 1956(1957), Ins. Micronesia 12: 70 (o*, 9, L*); Lien 1962,
Pacif. Ins. 4: 632 (distribution); Safyanova ef al. 1964, Zool. Zhur.
43: 1179 (distribution); Delfinado 1966, Mem. Amer. ent. Inst. 7:
122 (¢ *, 9, L*, P*).
Culex (Culiciomyia) pullus of Bohart and Ingram 1946, Navmed 1055: 31
(misidentification).
The adult male and female are recognized by the presence of a
prominent, black integumental spot on the upper meSepimeron. The male
terminalia may be distinguished by the absence of an enlarged hook at the
apex of the distimere, the presence of a prominent sternal accessory apical
Figure 42. C. (Culiciomyia) fragilis. Adult habitus: A, lateral aspect of
the male head; B, lateral aspect of the female head and thorax;
— C, dorsal aspect of the female wing; D, anterior surface of the
female legs; E, dorsal aspect of the female scutum and
scutellum; F, dorsal aspect of the female abdomen.
134 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.43
—
ean
C. (Culiciomyia) fragilis
Bram: Genus Culex in Thailand 135
lobe on the basimere, and by the number and shape of the setae on the sub-
apical lobe of the basimere. A unique feature of the fourth stage larva is the
presence of a lightly sclerotized ring slightly beyond the middle of the siphon,
giving the appearance of a false joint.
FEMALE. A small to medium sized species with distinctive colora-
tional features. Head. (Figure 45B). Proboscis and palpus uniformly dark
scaled; decumbent scales of the vertex light brown on the occiput, becoming
lighter until they are creamy white at the orbital line; erect scales yellowish
brown throughout. Thorax. (Figure 45B,E). Integument of the pleuron
light brown, with a distinct, velvety black spot present on the upper mesep-
imeron, anda lighter, but dark brown area stretching to the upper sterno-
pleuron; 1 strong lower mesepimeral bristle present. Wing. (Figure 45C).
Legs. (Figure 45D). Anterior surface of the hind femur predominantly
brown, but occasionally with some variable and randomly scattered white
scales; fore and mid legs similar to the hind legs. Abdomen. (Figure 45F).
Terga dark brown, with broad, pale basal bands and narrow, but variable,
pale apical bands which may be absent in some specimens; sterna uniformly
pale.
MALE. Head, (Figure 45A). Proboscis with a median tuft of
numerous strong setae on the ventral surface. Terminalia. (Figure 46C).
Subapical lobe of the basimere well developed, the most basal rod not on the
main lobe but separated from the other 2 rods which are subequal in size and
shape; following the basal rods are 2 narrow striated leaflets, 3 strong,
straight accessory setae, and a broad leaf which does not exhibit striations;
beyond the subapical lobe is a prominent sternal accessory lobe which is
covered with numerous long, fine setae; distimere with an apical crest of 15
or more recurved spines on the convex surface, a small hook opposite the
claw, and a variable number of setae on the basal half; lateral plate of the
phallosome with a large, heavily sclerotized basal tooth and 4 or 5 evenly
spaced denticles.
PUPA. (Figure 46A, B).
LARVA. (Figure 47). Head. Antenna with a narrow, dark, basal
ring; hair 1-C filamentous, its length slightly greater than half the distance
between the bases of the pair; 5, 6-C with 3 or 4 branches, pectinate. Thorax.
Integument glabrous; hairs 1,2,3-P single, pectinate, 3-P only slightly
shorter and slenderer than 1,2-P; 4,7, 8-P bifid, pectinate; 5, 6-P single,
pectinate; 14-P single, simple. Abdomen. Integument glabrous; comb
consisting of from 35 to 45 fan-shaped scales arranged in 3 irregular rows;
siphon index variable, ranging from 9:1 to 11:1, beyond the middle is a false
joint due to lack of sclerotization in an irregular ring; 3 pairs of subventral
tufts inserted on the siphon, 2 basad and 1 distad of the false joint; individual
tufts single or bifid, their length equal to or less than the width of the siphon
at the point of insertion; pecten consisting of from 8 to 11 teeth, restricted
to the basal fifth or less of the siphon; individual pecten tooth with a sharply
pointed distal spine and from 5 to 8 sharp, lateral barbs.
Figure 43. C. (Culiciomyia) fragilis. A,B, dorsoventral aspects of the
. pupa; C, dorsal aspect of the male terminalia.
136 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.44
C. (Culiciomyia) fragilis
Bram: Genus Culex in Thailand 1ST
TYPE DATA. Holotype female of annulaia from Kuching, Sarawak,
Malaya in the British Museum.
DISTRIBUTION. This species was first reported in THAILAND from
Bangkok and near Chumphon by Barraud and Christophers (1931), and Causey
(1937) also collected nigropunctatus in Bangkok. Material has been examined
during this study from Chiang Mai, Chon Buri, Krung Thep, Lampang, Mae
Hong Son, Nakhon Ratchasima, Nakhon Si Thammarat, Nonthaburi, Prachuap
Khiri Khan, Ranong, Sava Buri, Satun, Songkhla, and Thon Buri. The species
has also been recorded from BORNEO, INDIA, CEYLON, MALAYA, SINGA-
PORE, SUMATRA, JAVA, PHILIPPINES, HAINAN ISLAND, RYUKYU RETTO,
PALAU ISLANDS, CAROLINE ISLANDS, and TAIWAN. Two male specimens
have been examined in the British Museum from Rangoon, BURMA** (XII 62,
P. F. Mattingly).
During this study 26 individual rearings were examined from Thailand
as well as 72 females, 58 males, and 58 larvae.
TAXONOMIC DISCUSSION. The dark markings of the pleuron and the
exceptional siphon of the fourth stage larva clearly distinguishes nigropunctatus
from any other species of the subgenus in Thailand. It is very similar to
pullus Theobald from New Guinea, Australia, and the South Pacific, but may be
distinguished most noticeably on the basis of the male terminalia which are
quite distinct.
BIOLOGY. According to Bohart (1956(1957)), larvae are found
principally in or near rice fields in grassy ground pools in the Oriental region.
- Lien (1962) reported collecting larvae in Taiwan from hoof-marks, small
ground pools, and seepage pools with turbid water and grassy margins. How-
ever, Safyanova ef al. (1964) collected larvae of nigropunctatus only from
swamps in tropical rain forests in the Fucoy region of North Vietnam. Causey
(1937) collected larvae of this species in Thailand from small shady pools and
in water receptacles and collected adults at light traps throughout the year.
During this study larvae have been collected in Thailand from rock holes,
small ponds, puddles, rice fields, elephant tracks, buffalo hoof prints, a
coconut shell, various ditches, and once from a bamboo internode. Host
preference of the adult female, according to Colless (1959b), is birds, but ox
may Serve as a secondary host. This species has never been reported biting
man.
CULEX (CULICIOMYIA) PALLIDOTHORAX THEOBALD 1905
(Figures 48, 49, and 50)
Culex pallidothorax Theobald 1905, J. econ. Biol. 1: 32 (¢,9).
Culex albopleuva Theobald 1907, Mon. Cul. 4: 456 (¢); Edwards 1913, Bull.
ent. Res. 4: 235 (synonymy).
Culiciomyia annuloabdominalis Theobald 1910, Mon. Cul. 5: 236 (c'*, ?*);
Edwards 1913, Bull. ent. Res. 4: 235 (synonymy).
Culiciomyia pallidothorax (Theobald): Borel 1926, Arch. Insts. Pasteur
Indo-Chine 3-4: 30 (o*, 2, L*).
Figure 44. C. (Culiciomyia) fragilis. Fourth stage larva; dorsoventral
view of the head, thorax and abdomen, and lateral aspect of
— the terminal abdominal segments.
138 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
= y Yj Yj Yi
= y Z
: = we |) Y
: ES Y
—— Y
= —_ Y VY
——— 1g Y g
=—s Yy
= Y ys,
f/f] } Ih
Y
7,
ai = SEN Ne a < Wy
pat oo SRSA
Rites on s eis < : AX)
ne ae AN \
AW
1.0
cE
Mm. Misaki
C.(Culiciomyia) nigropunctatus
Bram: Genus Culex in Thailand 139
Culex (Culiciomyia) pallidothorax Theobald: Edwards 1932, in Wytsman,
Genera Insect. fasc. 194: 199 (taxonomy); Barraud 1934, Fauna Brit.
India, Diptera 5: 381 (o*,?, L); Bohart and Ingram 1946, Navmed
1055: 75 (o«*, 9*, L*); LaCasse and Yamaguti 1950, Mosq. Fauna
Japan and Korea: 182 (o*, 9*, L*, P*); Bick 1951, Pacif. Sci. 5: 419
(ecology); Hara 1957, Jap. J. exp. Med. 27: 52 (?*); Lien 1962,
Pacif. Ins. 4: 632 (distribution); Delfinado 1966, Mem. Amer. ent.
Inst. 7: 119 (L*, P).
The adult female can be separated from other members of the sub-
genus by the pale basal banding of the abdominal terga and by the dark patterns
on the integument of the pleuron. The adult male possesses a crest of spines
on the convex surface of the distimere of the terminalia and lacks a broad
leaf-like seta on the subapical lobe of the basimere.
FEMALE. A small to moderate sized species. Head. (Figure 48B).
Proboscis and palpus uniformly dark scaled; decumbent scales of the vertex
light brown at the occiput and becoming lighter towards the orbital line;
erect scales dark brown. Thorax. (Figure 48B,E). Integument of the pleuron
light brown, frequently tinged with green, with a distinctly darker brown pat-
tern which stretches from the prespiracular area across the prealar area and
terminates at the upper meSepimeron; another darker brown pattern present
on the upper sternopleuron; 1 or 2 strong lower mesepimeral bristles present.
Wing. (Figure 48C). Legs. (Figure 49D). All legs uniformly dark brown,
but occasionally with pale scaling on the ventral margin of the anterior surface
of the hind femur. Abdomen. (Figure 48F). Terga with rather broad,
usually convex, pale basal bands; sterna uniformly pale.
MALE. Head. (Figure 48A). Proboscis with a median tuft of setae
on the ventral surface. Terminalia. (Figure 49C). Subapical lobe of the
basimere well developed, the 3 rods closely placed, the basal rod shorter
and slenderer than the other 2 rods; additional strong setae and a narrow,
striated, leaf-like seta follows the rods; beyond the subapical lobe is a
prominent sternal accessory lobe which is covered with numerous long, fine
setae; distimere with a prominent crest of 5 or more recurved spines on
the convex surface and a distinct hook opposite the claw; lateral plate of the
phallosome with a prominent basal tooth and approximately 5 evenly spaced
denticles.
PUPA. (Figure 49A,B).
LARVA. (Figure 50). Head. Antenna with a narrow, dark basal ring;
head hair 1-C filamentous, its length slightly greater than half the distance
between the bases of the pair; 4-C single, simple; 5, 6-C with 2 or 3 branches,
pectinate. Thorax. Integument superficially glabrous, but when examined
critically, with indications of minute spicules (not illustrated); hairs 1,3-P
bifid, pectinate, 1-P slightly longer; 2-P single, pectinate; 4,7, 8-P bifid,
pectinate; 5,6-P single, pectinate; 14-P single, simple. Abdomen. Integu-
Figure 45. C. (Culiciomyia) nigropunctatus. Adult habitus: A, lateral
aspect of the male head; B, lateral aspect of the female head
ills and thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
140 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
“J, L. HWANG
C.(Culiciomyia) nigropunctatus
Bram: Genus Culex in Thailand ~141
ment similar to that of the thorax; comb consisting of from 25 to 40 fan-
shaped scales arranged in a triangular patch; siphon index variable, ranging
from approximately 4:1 to 6:1, the siphon greatly expanded medially; 4 pairs
of 3 or 4 branched subventral tufts inserted in a line beyond the pecten; the
length of the individual tufts less than half the width of the siphon at the point
of insertion; pecten consisting of from 7 to 10 teeth, restricted to the basal
third or less of the siphon; individual pecten tooth with a strong distal spine
and from 2 to 4 strong lateral barbs.
TYPE DATA. Lectotype of pallidothorax hereby designated: syntype
male (terminalia mounted and attached to pin), 'B.M. 1907-29, India, Capt.
James", in the British Museum. Holotype female (abdomen missing) of
albopleura from "India" in the British Museum. The type locality of
annuloabdominalis is Peradeniya and Hakgala, Ceylon, but the type specimen
is non-existent.
DISTRIBUTION. Edwards (1922) first recorded pallidothorax from
THAILAND. Subsequently, Barraud and Christophers (1931) reported the
species from Doi Suthep, and Causey (1937) recorded it from Bangkok,
"Chieng Moeang", and ''Scoed''. During this study collections of pallidothorax
have been made from: Chiang Mai, Chon Buri, Kanchanaburi, Lampang,
Lamphun, Mae Hong Son, Nakhon Nayok, Nakhon Ratchasima, Nan, Prachuap
Khiri Khan, Phare, Ranong, Sara Buri, and Tak. The species has also been
recorded from CEYLON, NEPAL, BURMA, INDOCHINA, MALAYA, CHINA,
TAIWAN, RYUKYU-RETTO, JAPAN, MOLUCCAS, NEW GUINEA, and the
PHILIPPINES.
Fifty individual rearings were examined from Thailand as well as 157
females, 165 males and 347 larvae.
TAXONOMIC DISCUSSION. Although quite distinctive, pallidothorax
has its closest affinity to thurmanorum and barrinus. Differences in the setae
on the subapical lobe of the basimere in the male terminalia, and larval
chaetotaxy and shape and color of the siphon clearly separate the species.
BIOLOGY. In Thailand, pallidothorax has been collected during this
study from puddles, rock holes, sumps, stream rock pools, rice fields,
elephant hoof prints, tin cans, and jungle streams from January to May,
August, September, and November. Barraud (1934) reported the habitat of
this species as tree holes, bamboo stems, shallow wells, stream and rock
pools, and swampy ground pools. Bohart and Ingram (1946) collected larvae
on Okinawa only from artificial containers, polluted ground pools, and rarely
in rock pools. Bick (1951) found larvae most frequently in puddles, followed
in frequency by artificial containers, pot holes, borrow pits, ditches,
Swamps, running creeks, coconut shells, tree holes, rock holes, and spathes
and leaves. He further stated that 77 percent of the collections were made
from temporary water, 67 percent of the habitats were exposed to sun, and
49 percent of the habitats demonstrated clear water in contrast to muddy,
stagnant, or polluted water. Lien (1962) recorded the larvae in Taiwan from
stream pools, rock pools, bamboo stumps, and artificial containers below
200 meters.
Figure 46. C. (Culiciomyia) nigropunctatus. A,B, dorsoventral aspects
of the pupa; C, dorsal aspect of the male terminalia.
C. (Culiciomyia) nigropunctatus
Bram: Genus Culex in Thailand 143
Host preference of the adult females is unknown, but apparently they
do not normally feed upon man (Feng 1938; Bohart and Ingram 1946; LaCasse
and Yamaguti 1950). However, Scanlon and Esah (1965) reported that small
numbers were collected biting and landing on man at the 2, 500 to 4, 500 feet
level on Pui Doi Mountain, Chiang Mai, Thailand. Iyengar (1938) reported
pallidothorax as a carrier of Wuchereria malayi. Hu (1940) induced starved
females to engorge on a human filarial patient and subsequent dissection
revealed the presence of infective Wuchereria bancrofti larvae, but he
concluded that pallidothorax did not play an important part in transmission.
CULEX (CULICIOMYIA) PAPUENSIS (TAYLOR) 1914
(Figures 51, 52, and 53)
Melanoconion papuensis Taylor 1914, Trans. R. ent. Soc. Lond. 1914: 201
(2).
Culex (Culiciomyia) papuensis (Taylor): Edwards 1924, Bull. ent. Res. 14:
397 (*, 2); King and Hoogstraal 1946, Proc. biol. Soc. Wash. 59:
146 (o'*, L*); Penn 1947, Pacif. Sci. 3: 73 (P*); Laird 1955, Bull.
ent. Res. 46: 286 (L*); Belkin 1962, Mosq. S. Pacif. I: 230 (c*, 9,
L*, P*); Assem and Bonne-Wepster 1964, Zool. Bijd. 6: 115 (2, L*);
Delfinado 1966, Mem. Amer. ent. Inst. 7: 120 (o*, 2, L*, P).
The adult female is poorly characterized but may be separated from
some species of the subgenus by the absence of pale abdominal banding, and
by the weak dorsocentral bristles. In the adult male, the distimere of the
terminalia and the structure and number of denticles on the lateral plate of
the phallosome are characteristic. The fourth stage larva may be recognized
by the robust head hair 1-C and the shape of the siphon.
FEMALE. A moderate to large species without distinctive color pat-
terns. Head. (Figure 51B). Proboscis and palpus uniformly dark scaled,
palpal segment IV over 3 times as long as III; decumbent scales of the vertex
yellowish brown on the occiput, becoming progressively lighter towards the
orbital line; erect scales dark brown. Thorax. (Figure 51B,E). Integument
of the pleuron light brown, frequently tinged with a yellowish green cast; a
faint, variable integumental pattern of brown is sometimes present on the
upper sternopleuron; 1 lower mesepimeral bristle present. Wing. (Figure
91C). Legs. (Figure 51D). Anterior surface of hind and mid femora dark,
but occasionally with a stripe of pale scales on the ventral margin; fore
femur and all tibiae and tarsi uniformly dark, although the fore femur may
occasionally have some pale scales on the ventral margin. Abdomen.
(Figure 51F). Terga uniformly dark; sterna pale.
MALE. Head. (Figure 5A), Proboscis with a median tuft of strong
setae inserted on the ventral surface. Terminalia. (Figure 52C). Subapical
lobe of the basimere moderately developed, the basal rod shorter than, and
separated from the other 2 rods which are subequal in strength and length;
Figure 47. C. (Culiciomyia) nigropunctatus. Fourth stage larva: dorso-
ventral view of the head, thorax and abdomen, and lateral
iia aspect of the terminal abdominal segments.
144 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
ODORANT AIAN
C. (Culiciomyia) pallidothorax
Bram: Genus Culex in Thailand 145
beyond the rods are a Series of 3 strong, broad, straight setae and 5 or more
accessory setae, but no leaf-shaped seta; beyond the subapical lobe is a
prominent sternal accessory lobe which is covered with numerous long, fine
setae; distimere with a crest of 3 or 4 recurved spines on the convex surface
and with a small hook opposite the claw; lateral plate of the phallosome with
a prominent, heavily sclerotized basal tooth and from 4 to 8 evenly spaced
denticles.
PUPA. (Figure 52A,B).
LARVA. (Figure 53). Head. Antenna with a narrow dark basal ring;
head hair 1-C robust, lightly pigmented, frequently possessing accessory
lateral spicules, its length somewhat greater than half the distance between
the bases of the pair; 4-C single, simple; 5-C usually trifid, pectinate; 6-C
usually with 4 branches, pectinate. Thorax. Integument with extremely
minute spicules arranged in an irregular pattern, particularly on the antero-
lateral margin (not illustrated); hair 1-P trifid, pectinate; 2-P single, pecti-
nate, shorter than 1-P; 3-P trifid, pectinate, shorter than 1-P; 4-P bifid,
pectinate; 5,6-P single, pectinate; 7-P trifid, pectinate; 8-P with 4 pectinate
branches; 14-P single, simple. Abdomen. Integument glabrous; comb
consisting of approximately 30 to 40 fan-shaped scales arranged in a broad,
triangular patch; siphon index variable, ranging from 4.5:1 to 5.5:1, the
siphon greatly expanded medially; 4 pairs of subventral tufts inserted in a
line beyond the pecten; individual tufts with from 6 to 10 branches, their
length considerably less than the width of the siphon at the point of insertion;
pecten consisting of from 3 to 5 teeth, restricted to approximately the basal
third of the siphon; individual pecten tooth with a strong distal spine and 3 or
4 smaller lateral barbs.
TYPE DATA. Holotype female of papuensis from Lahekamu Gold
Field, Papua, New Guinea in the School of Public Health and Tropical Medi-
cine, University of Sydney, Australia.
DISTRIBUTION. In THAILAND**, this species has been collected
from: Nakhon Ratchasima, Narathiwat, Phattalung, and Songkhia. Previous
records have been from NEW GUINEA, BISMARCK ARCHIPELAGO, the
SOLOMON ISLANDS, the PHILIPPINES, and 2 larvae in the British Museum
have been examined from "Java or Sumatra?, Tjoeroel".
During this study 84 individual rearings were examined as well as 17
females, 15 males, and 99 larvae.
TAXONOMIC DISCUSSION. This species shows no particularly close
affinity to any of the other species of the subgenus in Thailand, although no
outstandingly distinctive features may be observed in the adult female. In the
male, the terminalia is similar to nigropunctatus and pallidothorax, but the
setae on the subapical lobe of the basimere are distinctive, the terga lack pale,
Figure 48. C. (Culiciomyia) pallidothorax. Adult habitus: A, lateral aspect
of the male head; B, lateral aspect of the female head and
—— thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
146 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.49
} iy PP his
Aon ett
iy TG eT Zak
a ee ee
Cur
(lh
AI HQ
vee,
i 1)
SS
C. (Culiciomyia) pallidothorax
Bram: Genus Culex in Thailand 147
basal bands, and dark integumental markings are absent from the pleuron.
In the fourth stage larva, the siphon is similar to that of pallidothorax but -
the distinctive head hair 1-C and other features of the chaetotaxy are
diagnostic. ‘
BIOLOGY. In Thailand, larvae have been collected from stream pools,
rock pools, elephant tracks, and a large root hole. Assem and Bonne-
Wepster (1964) reported collecting larvae in New Guinea from ground water
and container habitats in shade or half shade, and in either clear water or
water filled with leaves; sometimes they were present in putrid water found
in hollowed sage-palm trunks. Also in New Guinea, Bick (1951) collected
larvae most frequently from artificial containers, followed by puddles, pot
holes, borrow pits, ditches, rock holes, and tree holes. Usually vegetation
was absent from the habitats, but it was 1 of the few species which Bick col-
lected frequently in both surface water and container habitats. King and
Hoogstraal (1946) found larvae in putrid water in hollowed sage trunks, in
wooden kegs, tin food containers, tree holes, and drums. In the Solomon
Islands larvae were collected by Belkin (1962) from large tree holes,
artificial containers, and ground pools. Adult host preference and bionomics
are unknown, but man has not been reported as a host.
CULEX (CULICIOMYIA) SCANLONI, N. SP.
(Figures 54, 55, and 56)
The adult female cannot be distinguished with certainty from fragilis.
The adult male may be recognized by the setae on the subapical lobe of the
basimere and by structures on the lateral plate of the phallosome. The
fourth stage larva is characterized by the shape and length of the siphon and
by the shape of the pecten tooth.
FEMALE. A small species without striking colorational features.
Head. (Figure 54B). Proboscis and palpus uniformly light brown with a
grayish cast; decumbent scales of the vertex creamy white; erect scales dark
brown. Thorax. (Figure 54B,E). Integument of the pleuron yellowish brown,
frequently tinged with green in some specimens, without consistent dark
patterns present; 1 lower mesepimeral bristle present. Wing. (Figure 54F).
Legs. (Figure 54D). Anterior surface of the femora grayish brown, with
variable white stripes on the ventral margins; tibiae and tarsi brown, but ap-
pearing grayish when struck by light at an oblique angle. Addomen. (Figure
54F). Terga brown; sterna pale.
MALE. Head, (Figure 54A). Proboscis with a median tuft of from
2 to 4 strong setae on the ventral surface. Terminalia. (Figure 55C). Sub-
apical lobe of the basimere well developed, but rather narrow; the basal rod
Separated from the other 2 rods, beyond which are 2 sigmoid and 1 straight
accessory setae and a broad, striated leaf; distimere with a crest of 15 or
more recurved spines which are absent only from the extreme base and apex
of the convex surface, with a small hook opposite the claw, and with several
A ees
Figure 49. C. (Culiciomyia) pallidothorax. A,B, dorsoventral aspects
<< of the pupa; C, dorsal aspect of the male terminalia.
148 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C.(Culiciomyia) pallidothorax
Bram: Genus Culex in Thailand 149
fine setae on the basal half; lateral plate of the phallosome with a rather large,
heavily sclerotized basal tooth and approximately 6 sharply pointed denticles.
PUPA. (Figure 55A, B).
LARVA. (Figure 56). Head. Antenna with a narrow, dark basal ring;
head hair 1-C filamentous, its length slightly greater than half the distance
between the bases of the pair; 4-C single, simple; 5-C trifid, pectinate; 6-C
bifid, pectinate. Thovax. Integument superficially glabrous, but critical
examination reveals the presence of minute spicules (not illustrated); hairs
1,2,3-P single, pectinate, 3-P shorter and slenderer than 1, 2-P; 4,7, 8-P
bifid, pectinate; 5,6-P single, pectinate; 14-P single, simple. Abdomen.
Integument similar to that of the thorax; comb consisting of approximately 35
to 45 fan-shaped scales arranged in a broad, triangular patch; siphon index
variable, ranging from 9.5:1 to 10.5:1, the siphon gently, but characteristically
curved; 3 pairs of subventral tufts inserted beyond the pecten; individual tufts
single or bifid, their length approximately equal to the width of the siphon at
the point of insertion; pecten consisting of approximately 15 teeth restricted
to the basal fifth or less of the siphon; individual pecten tooth with a fine
distal spine and 1 or 2 fine lateral barbs as well as a single, strong, stout,
bluntly rounded basal barb.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia slide-mounted) from Quan Ga Lang, Satun, Thailand, 12 March
1965, E. L. Peyton, from a small, lightly shaded rock pool, ST 5-10,
deposited in the U. S. National Museum, No. 69348. Paratype female with
associated larval and pupal skins slide mounted from Huey Yang Water fall,
Prachuap Khiri Khan, Thailand, 27 March 1964, K. Mongkolpanya, from a
rock pool, PR 159-30, deposited in the U. S. National Museum, No. 69348.
Paratype male with associated larval and pupal skins and terminalia slide-
mounted from 26 km. Ngao to Payae Road, Lampang, Thailand, 11 June
1964, E. L. Peyton, from a roadside puddle, LG 8-30, deposited in the U. S.
National Museum, No. 69348.
This species is named in honor of Lt. Col. John E. Scanlon, MSC,
USA in recognition of his contributions to Medical Entomology in Thailand.
DISTRIBUTION. In THAILAND, scanloni has been collected from:
Chon Buri, Kanchanaburi, Lampang, Mae Hong Son, Nakhon Ratchasima,
Narathiwat, Prachuap Khiri Khan, Ranong, Satun, and Surat Thani.
Larvae of this species have been examined from Subic Bay, and San
Jose, Mindoro, PHILIPPINE ISLANDS, Kota Belud, NORTH BORNEO, and
Bien Hoa, SOUTH VIETNAM. During this study 15 individual rearings have
been examined as well as 11 females, 9 males, and 61 larvae.
TAXONOMIC DISCUSSION. This species demonstrates extremely
close affinity to fragilis, and in the adult stage the 2 species are virtually
inseparable. In the fourth stage larvae, however, distinct differences exist
in the chaetotaxy of the head (eg. 5-C trifid, 6-C bifid in scanloni, 5, 6-C
with from 5 to 8 branches in fragilis), prothorax (eg. 14-P single in scanloni,
bifid in fragilis), but particularly in the length and shape of the siphon
which is very long and gently curved in scanloni, and in the shape of the pecten
LL AL LL CL LL TL EL pe eeenpungenengyenersnenerenesenngums
Figure 50. C. (Culiciomyia) pallidothorax. Fourth stage larva: dorso-
< ventral view of the head, thorax and abdomen, and lateral
aspect of the terminal abdominal segments.
150 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fr ig.5d
C. (Culiciomyia ) papuensis
Bram: Genus Culex in Thailand 151
tooth.
BIOLOGY. Larvae of scanloni have been collected from rock pools,
stream pools, puddles, and elephant hoof prints. Nothing is known of the
adult habits or biology.
CULEX (CULICIOMYIA) SPATHIFURCA (EDWARDS) 1915
(Figures 57, 58, and 59)
Culiciomyia spathifurca Edwards 1915, Bull. ent. Res. 5: 284 (¢).
Culex (Culiciomyia) spahtifurca (Edwards): Edwards 1922, Indian J. med.
Res. 10: 472 (distribution); Edwards 1926, Bull. ent. Res. 17: 120
(L); Delfinado 1966, Mem. Amer. ent. Inst. 7: 123 (o*, 92, L, P).
Culex (Culiciomyia) stylifurcatus Carter and Wijesundara 1948, Ceylon J.
Sci. (B) 23: 145 (o*, 2, L*); Mattingly 1955, zz Iyengar and Menon,
Bull. ent. Res. 46: 10 (synonymy).
The adult female cannot be separated with certainty from several other
members of the subgenus. In the adult male, the terminalia is unique in
possessing a bifurcate distimere. The fourth stage larva may be recognized
by having thoracic hairs 1,2,3-P single, siphon index approximately 5:1, head
hairs 5,6-C with 3 or 4 branches, and the subventral tufts of the siphon
single.
FEMALE. A small to medium sized species without outstanding
colorational or anatomical features. Head. (Figure 57B). Proboscis and
palpus uniformly dark scaled; decumbent scales of the vertex yellowish brown
at the occiput, becoming lighter at the orbital line; erect scales yellowish
brown. Thorax. (Figure 57B,E). Integument of the pleuron light brown,
occasionally faintly tinged with green and with a slightly darker integumental
spot sometimes present on the postspiracular plate; 1 lower mesepimeral
bristle present. Wing. (Figure 57C). Legs. (Figure 57D). Anterior
surface of the hind femur dark brown, with a variable but distinct stripe of
pale scales on the ventral margin; hind tibia also with some pale scales on
the ventral margin; hind tarsus as well as the fore leg and mid tibia and
tarsus dark brown; mid femur similar to the hind femur. Abdomen. (Figure
57F). Terga uniformly dark; sterna pale.
MALE. Head. (Figure 57A). Proboscis with a median tuft of strong
setae inserted on the ventral surface. Terminalia. (Figure 58C). Basimere
with the subapical lobe well developed; basal rod slender, separated from the
2 other rods which are subequal in length and inserted on a distinct tubercle;
beyond the basal rods are 2 foliaceous setae and a broad leaf, as well as a
number of long, straight accessory setae; distimere furcate, the proximal
arm smooth and sigmoid, the distal arm broad, with numerous denticles on
the outer margin, and with a variable number of short setae; lateral plate of
Figure 51. C. (Culiciomyia) papuensis. Adult habitus: A, lateral aspect
of the male head; B, lateral aspect of the female head and
~<— thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
152 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
rig.G72
2
> 9h
JIN 4
“JL. HWANG
B
Bram: Genus Culex in Thailand 153
the phallosome rather small, with a basal tooth and approximately 7 denticles.
PUPA. (Figure 58A, B).
LARVA. (Figure 59). Head. Antenna with a narrow, dark basal
ring; head hair 1-C filamentous, its length slightly less than half the distance
between the bases of the pair; 4-C single, simple; 5, 6-C with three pectinate
branches. Thorax. Integument glabrous; hairs 1,2,3-P single, pectinate,
3-P shorter and slenderer than 1,2-P; 4,7, 8-P bifid, pectinate; 5,6-P
single, pectinate; 14-P single, simple. Abdomen. Integument glabrous;
comb consisting of approximately 40 to 50 fan-shaped scales arranged in a
triangular patch; siphon index variable, approximately 5:1 to 6:1; 3 pairs of
subventral tufts inserted in a line beyond the pecten; individual tufts
represented by a single seta whose length is considerably less than the width
of the siphon at the point of insertion; pecten consisting of approximately 11
teeth restricted to the basal 1/5 of the siphon; individual pecten tooth with a
strong apical spine and 4 or 5 strong lateral barbs.
TYPE DATA. Holotype male of spathifurca (terminalia mounted and
attached to pin) from Kuching, Sarawak, Borneo in the British Museum.
Holotype male of stylifurcatus (terminalia slide-mounted, but both basimeres
and distimeres missing) from Ratmalana and Palwatte Ganga River at
Dinipitiya, Ceylon in the British Museum.
DISTRIBUTION. Causey (1937) first recorded spathifurca in THAI-
LAND from the vicinity of Bangkok. During this study specimens have been
identified from Chiang Mai, Chon Buri, Lampang, and Ranong. The species
has also been recorded from BORNEO, JAVA, SINGAPORE, MALAYA,
CEYLON, MALDIVE ISLANDS, TAIWAN, NEW GUINEA, and the PHILIPPINE
ISLANDS.
Three individual rearings have been studied from Thailand as well as
15 females, 12 males, and 14 larvae.
TAXONOMIC DISCUSSION. Although the adult female cannot be
separated with certainty from fragilis and scanloni, the male terminalia
exhibits unique features in the bifurcate distimere, the shape and number of
structures on the subapical lobe of the basimere, and by the shape of the
phallosome. The fourth stage larva shows some affinity to fragilis but may
be separated by the chaetotaxy of the head and siphon.
BIOLOGY. Causey (1937) reported that the larvae of spathifurca were
collected from all types of artificial receptacles, temporary pools, canals,
rice fields, and swamps of brackish water in the vicinity of Bangkok. Larvae
were collected by Edwards (1926) from pots of urine in Singapore, and Carter
and Wijesundara (1948) collected larvae as well as adults from crab holes in
Ceylon. During this study larvae were collected in Thailand from elephant
tracks, puddles, brackish water in nipa palm leaves, rock pools, ditches,
and a bamboo stump.
The normal host of the adult female is apparently birds, but
precipitin tests showed that ox may also serve as a host (Colless 1959).
Slooff and van Dijk (1961) reported that on Pam Island, Netherlands New
Guinea, spathifurca constituted 3 percent of the total number of mosquitoes
captured during night catches on human bait. They further stated that 7 1/2
re rene i tin i es ileal ac Ui a ge
Figure 52. C. (Culiciomyia) papuensis. A,B, dorsoventral aspects of
<< the pupa; C, dorsal aspect of the male terminalia.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
154
Fig.53
aS
Po
Aff
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OY)
ih
y
“HZ,
a”
SAY
N
NN
SS
SOL... HWANG
\\\
C. (Culiciomyia) papuensis
Bram: Genus Culex in Thailand 155
days after feeding on a human carrier of Wuchereria bancrofti 1 specimen of
spathifurca was found to have 4 normally developed second instar larval W.
bancrofti in the thorax, and they concluded that spathifurca is a potential
vector of filariasis on Pam Island.
CULEX (CULICIOMYIA) SPICULOTHORAX,-N. SP.
(Figure 60)
The fourth stage larva is characterized primarily by the nature of the
thoracic spicules, but other characteristics which are diagnostic include the
pectinate head hair 4-C, the spiculate clypeus, and the shape and chaetotaxy
of the siphon.
FEMALE, MALE, PUPA. Unknown.
LARVA. (Figure 60). Head. Antenna with a narrow dark basal ring;
clypeus with an irregular pattern of minute spicules; head hair 1-C filamentous,
its length slightly greater than the distance between the bases of the pair; 4-C
single, pectinate; 5,6-C with 3 or 4 pectinate branches. Thovax. Integument
conspicuously spiculate, the spicules short, conical, and dark; hairs 1,3-P
bifid, pectinate; 2-P single, pectinate; 4,7,8-P bifid, pectinate; 5,6-P single,
pectinate; 14-P single, pectinate. Abdomen. Integument spiculate, the
Spicules not nearly as prominent as those of the thorax; comb consisting of
approximately 35 to 50 fan shaped scales arranged in a triangular patch of 3
irregular rows; siphon index variable, ranging from 3.5:1 to 4:1; 4 pairs of
subventral tufts inserted in a line beyond the pecten; individual tufts with
from 3 to 5 branches, their length variable, usually slightly greater than the
width of the siphon at the point of insertion; pecten consisting of from 10 to
15 teeth restricted to the basal third of the siphon or less; individual pecten
tooth with a strong distal spine and 3 strong, lateral barbs.
TYPE DATA. Holotype larva from Ban Khum Klang, Chiang Mai,
Thailand, 19 November 1963, J. E. Scanlon, ex. bamboo stump, CM 212,
deposited in the U. S. National Museum, No. 69349. Six paratype larvae
with the same data as the holotype, also deposited in the U. S. National Mu-
seum.
DISTRIBUTION. That of the type locality, Ban Khun Klang, Chiang
Mai, THAILAND. This species is known only from the type series.
TAXONOMIC DISCUSSION. Although the adults are unknown, the
fourth stage larva is sufficiently distinct to warrant description at this time.
The distinctive thoracic and abdominal spiculation and features of the chae-
totaxy easily separate spiculosus from other known members of the sub-
genus.
BIOLOGY. Other than data from the type habitat, nothing is known of
the biology or habits of this species.
Figure 53. C. (Culiciomyia) papuensis. Fourth stage larva: dorsoventral
< view of the head, thorax and abdomen, and lateral aspect of
the terminal abdominal segments.
156 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C.(Culiciomyia) scanloni
Bram: Genus Culex in Thailand 157
CULEX (CULICIOMYIA) TERMI D. THURMAN 1955
(Figures 61, 62, and 63)
Culex (Culiciomyia) termi D. Thurman 1955, Proc. ent. Soc. Wash. 57:
18 (o*, 2, P, L*).
The adult female can be distinguished from other members of the
subgenus by the overall pale scaling, the light dorsal wing scales, and the
broad apical banding of the abdominal terga. The male terminalia exhibits
a lightly sclerotized phallosome with denticles reduced in size, anda
distinctive number and arrangement of setae on the subapical lobe of the
basimere. The fourth stage larva is unique in having the siphon index 30:1
or greater, thoracic hair 4-P with from 6 to 8 branches, and the comb
consisting of approximately 70 scales.
FEMALE. In general, a moderately sized species with an overall
pattern of light scales. Head. (Figure 61B). Proboscis light brown to
white medially, but somewhat darker proximally and distally; palpus with
dark brown scales on the apical and subapical segments, reddish-brown
scales basally, with isolated light scales also present; decumbent scales of
the vertex uniformly creamy white; erect scales yellowish-white throughout
except for a few dark scales on the posterior margin. Thorax. (Figure
61B, E). Scutum and scutellum yellowish with a uniform pattern of pale
scales; integument of the pleuron uniformly light brown, with a variable dark
stripe running from beneath the anterior spiracle to the upper mesepimeron;
1 lower mesSepimeral bristle present and with a small, variable patch of
pale scales on the upper sternopleuron. Wing. (Figure 61C). Dorsal wing
scales predominantly pale. Legs. (Figure 61D). Femora and tibiae pre-
dominantly pale with some dark scales on the dorsal margins; tarsi dark.
Abdomen. (Figure 61F). Terga with apical pale bands that cover the greater
part of the segments; sterna completely pale.
MALE. Head. (Figure 61A). Proboscis predominantly pale, with a
median tuft of strong bristles on the ventral and lateral surfaces; palpus pre-
dominantly pale with some dark scales on the apical and subapical segments.
Terminalia. (Figure 62C). Subapical lobe of the basimere well developed,
the 3 basal rods followed by 5 narrow leaflets; distimere smooth, with a hook
opposite the claw and with at least 1 basal seta; lateral plate of the phallo-
some lightly sclerotized, with a basal tooth and approximately 7 or more
very small, inconspicuous denticles.
PUPA. (Figure 62A,B). Significant in the absence of paddle hairs
ta
LARVA. (Figure 63). Head, Antenna with a narrow, dark basal
ring; head hair 1-C filamentous, its length slightly less than half the distance
between the bases of the pair; 4-C single, simple; 5, 6-C with 3 or 4 branches,
Figure 54. C. (Culiciomyia) scanloni. Adult habitus: A, lateral aspect
of the male head; B, lateral aspect of the female head and
thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdo-
men.
others
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
158
/
ia) scanion
¥.
icfomy
fs
C.(Cu
Bram: Genus Culex in Thailand 159
pectinate. Thorax. Integument covered with a dense pattern of sharply
pointed spicules; hairs 1,2-P single, pectinate; 3-P bifid, pectinate, its
length less than that of 1,2-P; 4-P with from 6 to 8 pectinate branches; 5, 6-P
single, pectinate; 7-P with 4 pectinate branches; 8-P trifid, pectinate; 14-P
single, simple. Abdomen. Integument similar to that of the thorax; comb
consisting of approximately 60 to 70 fan-shaped scales arranged in a tri-
angular patch; siphon index approximately 30:1 or more, the siphon slender
throughout its length; 6 pairs of subventral tufts widely spaced on the siphon
beyond the pecten; individual tufts single or bifid, their length slightly greater
than the width of the siphon at the point of insertion; pecten consisting of from
9 to 17 teeth; individual pecten tooth with a strong distal spine and approximate-
ly 9 fine lateral barbs.
TYPE DATA. Holotype male from Ngao, Lampang, Thailand in the U.
S. National Museum.
DISTRIBUTION. Known only from THAILAND, from Lampang and
Mae Hong Son. During this study the following specimens have been examined:
7 females, 5 males, and 105 larvae from the original Thurman collection,
and 42 females and 28 males, 41 with their associated larval and pupal skins,
and 35 larvae collected from Mae Hong Son.
TAXONOMIC DISCUSSION. The pale adult habitus and the exceptionally
long larval siphon unquestionably separate termi from all other species of the
subgenus in Thailand.
BIOLOGY. Larvae of termi were collected by D. Thurman (1955)
from elephant-track depressions in a marsh overgrown with luxuriant vegeta-
tion where the water was highly polluted with elephant dung and filled with
brown algae; collections were made during March, July, August, and Novem-
ber. The 7 collections from Mae Hong Son were all from elephant hoof prints
in a Secondary rain forest at 600 m. elevation or more, and were made during
September. Nothing is known of the adult habits.
CULEX (CULICIOMYIA) THURMANORUM, N. SP.
(Figures 64 and 65)
Culex (Culiciomyia) viridiventer of E. Thurman 1959, Univ. Md. Agric. Exp.
Sta. Bull. A-100: 122 (misidentification); E. Thurman 1963, Proc.
IX Pacif. Sci. Cong. 9: 55 (misidentification).
The adult female may be recognized by the pale basal banding of the
abdominal terga and by the dark patterns on the integument of the pleuron.
The adult male exhibits banded abdominal terga, a crest at the apex of the
distimere, and lacks a leaf-like structure on the subapical lobe of the basi-
mere. The fourth stage larva is distinguished from all other members of
the subgenus by possessing prominent, stellate setae on both the thorax and
abdomen, and by the modified scalelike spicules of the abdominal integument.
FEMALE. Head. Proboscis and palpus uniformly dark scaled; de-
cumbent scales of the vertex light brown at the occiput and becoming almost
Figure 54, C. (Culiciomyia) scanloni. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax; C, dorsal
oe aspect of the female wing; D, anterior surface of the female legs; E,
dorsal aspect of the female scutum and scutellum; F, dorsal aspect of
the female abdomen.
160 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig. oo
C.(Culiciomyia) scanloni
Bram: Genus Culex in Thailand 161
white at the orbital line; erect scales dark brown. Thorax. Integument of
the pleuron light brown, with a dark area on the postspiracular plate and a
dark band stretching across the upper sternopleuron and upper mesepimeron
and usually with another brown pattern on the sternopleuron; 1 or 2 strong,
lower mesepimeral bristle present. Wing. All dorsal wing scales uniformly
light brown. Legs. Anterior surface of the hind femur pale, with a narrow
dark apical band that extends along the dorsal margin as a dark stripe; hind
tibia and tarsus as well as the fore legs uniformly dark brown. Abdomen.
Terga dark brown, with moderately broad pale basal bands on segments III-
VII; sterna uniformly pale.
MALE. Head. Proboscis with a ventromedian tuft of setae.
Terminalia. (Figure 64C). Subapical lobe of the basimere well developed,
with 4 strong rod-like setae, several accessory setae, and with a small, in-
conspicuous leaf-like structure; distimere with a crest of 6 or more small,
recurved spines on the convex surface and with a small hook opposite the
claw; lateral plate of the phallosome strongly sclerotized, with a prominent
basal tooth and 7 or more evenly spaced, short denticles.
PUPA. (Figure 64A,B).
LARVA. (Figure 65). Head, Hair 1-C is not filamentous, the com-
mon condition in the subgenus, nor is it grossly robust, but assumes an
intermediate form; 4-C simple, single or double; 5-C with 5 or 6 branches,
pectinate; 6-C with 4 or 5 branches, pectinate; the frontal head hairs placed
far forward. Thorax. Integument densely spiculose, with prominent,
sharply pointed spicules; hair 1-P bifid, pectinate; 2,3-P single, pectinate,
3-P considerably shorter and slenderer than 1, 2-P; 4-P bifid or trifid,
pectinate; 5,6-P single, pectinate; 7,8-P bifid, pectinate; 14-P single,
simple; 1-M, T with 5 or 6 stellate branches (this condition sharply contrasts
the normal condition in which 1-M, T are short and dendritic); 3,13-T also
stellate. Abdomen. Integument densely spiculose, the individual spicules
modified, forming a uniform pattern of scale-like platelets over the entire
abdominal integument; terga I through VI with at least 2 pairs of stellate
setae; comb consisting of approximately 30 scales arranged in a rather broad
triangular patch; individual scales elongate with the apical spines stronger
than the lateral ones; siphon index ranging from 7:1 to 8.5:1; 3 pairs of sub-
ventral tufts evenly spaced beyond the pecten; individual tufts consisting of a
single seta, their length slightly greater than the width of the siphon at the
point of insertion; pecten consisting of from 6 to 12 teeth restricted to less
than the basal half of the siphon, the distal teeth more widely spaced than the
proximal ones; individual pecten tooth with a prominent distal spine and 3
sharply pointed, strong lateral barbs. The stellate setae of the thorax and
abdomen are outstandingly distinctive; variation has been noted in the number
and width of the branches as well as the relative number of lateral barbs.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia slide-mounted) from Doi Chang, Mae Hong Son, Thailand, 12
September 1966, K. Mongkolpanya, from elephant hoofprints in a secondary
rain forest, at 600 m. elevation, deposited in the U. S. National Museum,
Figure 55. C,. (Culiciomyia) scanloni. A,B, dorsoventral aspects of
ioe the pupa; C, dorsal aspect of the male terminalia.
162 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig: 56
C. (Culiciomyia) scanloni
Bram: Genus Culex in Thailand 163
No. 69350. Paratypes: 3 males and 7 females with their associated larval
and pupal skins, and with the same data as the holotype deposited in the U. S.
National Museum. This species is named in honor of D. C. Thurman and
E. B. Thurman (Mrs. C. Schwartzwalder) in recognition of their contributions
to the understanding of the mosquito fauna of Thailand.
DISTRIBUTION. In addition to the type locality, thurmanorum has
also been collected from Ban Mae Ho Kae No Chae and Ban Mae Ho Nua, both
also in Mae Hong Son, THAILAND, and the specimens in the Thurman col-
lection are from the Ngao district of Lampang.
The following specimens were studied in addition to the type series:
29 females and 24 males, 5 with their associated larval and pupal skins, and
101 larvae. |
TAXONOMIC DISCUSSION. Although the adult female is extremely
similar to pallidothorax, the fourth stage larva is so distinctive that it cannot
be confused with any other species; the larva of vividiventery differs in many
points from thurmanorum including the absence of the distinctive stellate
setae. The male terminalia lacks the conspicuous leaf-like seta on the sub-
apical lobe of the basimere which is present in pallidothorax.
BIOLOGY. This species has been collected only in the immature
stages from secondary rain forests in mountainous areas (elevation 600 m. or
above). Of 23 different collections, all but 2 were from elephant hoof prints;
the other 2 collections were from a puddle and a stream margin near the
hoofprint environments of other collections. The collections from Lampang
were made in March, those from Mae Hong Son during September. Larvae
have been collected in association with termi, pallidothorax, bailyi, and
barrinus. Nothing is known of the habits of the adults.
SUBGENUS CULEX LINNAEUS 1758
Culex Linnaeus 1758, Systema Naturae, 10th ed. 1: 602. Type species:
Culex pipiens Linnaeus (selection of Latreille 1810).
Heteronycha Lynch Arribalzaga 1891, Rev. Mus. La Plata 1: 373, 2: 155.
Type species: Heteronycha dolosa Lynch Arribalzaga.
Lasioconops Theobald 1903, Mon. Cul. 3: 235. Type species: Lasioconops
poticilibes Theobald.
Heptaphlebomyia Theobald 1903, Mon. Cul. 3: 336. Type species:
Heptaphlebomyia simplex Theobald.
Pseudoheptaphlebomyia Ventrillon 1905, Bull. Mus. Hist. nat. Paris 11: 427.
Type species: Pseudoheptaphlebomyia madagascariensis Ventrillon.
Trichopronomyia Theobald 1905, Ann. hist.-nat. Mus. hung. 3: 98. Type
species: Trichopronomyia annulata Theobald (not Schrank 1776).
Aporoculex Theobald 1907, Mon. Cul. 4: 316. Type species: Aporoculex
punctipes Theobald.
Leucomyia Theobald 1907, Mon. Cul. 4: 372 (preoccupied by Brauer and
Bergenstamm 1891). Type species: Culex gelidus Theobald.
SSsSSnaE ERR RERIneReimmmieceacengeesnanne rapee meee RTT imme onc se TC PaO ne en
Figure 56. C. (Culiciomyia) scanloni. Fourth stage larva: dorsoventral
: view of the head, thorax and abdomen, and lateral aspect of
the terminal abdominal segments.
164 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C.(Culiciomyia/ spathifurca
Bram: Genus Culex in Thailand 165
Oculeomyia Theobald 1907, Mon. Cul. 4: 515. Type species: Oculeomyia
sarawaki Theobald.
Theobaldiomyia Brunetti 1912, Rec. Indian Mus. 4: 462. Type species:
Culex gelidus Theobald.
Phalangomyia Dyar and Knab 1914, Insec. Inscit. menst. 2:58. Type
species: Phalangomyia debilis Dyar and Knab.
Laiomyia Izquierdo 1916, Tesis. Col. Est. Puebla: 65. Type species:
Culex stigmatosoma Dyar.
Culex (Tvansculicia) Dyar 1917(1918), Insec. Inscit. menst. 5: 184. Type
species: Culex eleuthera Dyar.
Culex (Cacoculex) Dyar 1918, Insec. Inscit. menst. 6: 100. Type species:
Culex habilitator Dyar and Knab.
The adult female may be recognized by the presence of distinct scale
patches on the pleuron and by the presence of a median band of white scales
on the proboscis in all but the 3 species of the pipiens group. The adult male
is distinguished by the above characters as well as the complex phallosome
of the terminalia. The fourth stage larva exhibits a distinct pecten restricted
to the basal third or less of the siphon, 10 or more tufts in the ventral brush,
and thoracic hair 3-P single, of the same order of length and thickness as
1-P.
FEMALE. Medium to large sized species. Head. Proboscis with
or without a median dorsal band of pale scales (absent only in members of
the pipiens group). Thorax. Scales of the scutum very dense, smooth,
with or without a distinct pattern; acrostichal bristles well developed, as are
the anterior dorsocentral, posterior dorsocentral, supraalar, prescutellar,
and scutellar bristles; pleuron with distinct scale patches and with extensive,
but varied bristle patterns; 1 or 2 lower mesSepimeral bristles present in
those species without a banded proboscis, no lower mesepimeral bristles
present when proboscis is banded; wings with dense scaling which may be
either all dark or with dark and light scales; legs with or without pale bands.
Abdomen. Terga with or without basal and/or apical pale bands.
MALE. Head. Palpus with 5 segments and usually longer than the
proboscis, the distal 2 segments upturned and without distinctive lanceolate
scales on the ventral surface of segment III; antenna approximately as long as
the proboscis, without distinctive specialized hairs or scales. Terminalia.
Subapical lobe of the basimere well developed, having strongly developed
setae and always with at least 1 leaf-shaped seta; distimere with or without
distal annulations, somewhat expanded medially in a few species; phallosome
complex, the lateral plate frequently spiculose or rugulose, with both an
inner and outer division; paraproct crowned with a dense tuft of spicules;
cercal setae short, variable in number; basal sternal process always
present, but variable.
LARVA. Head. Hair 1-C variable in shape; 16,17-C absent. Thorax.
Figure 57. C. (Culiciomyia) spathifurca. Adult habitus: A, lateral aspect
of the male head; B, lateral aspect of the female head and
tif thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
166 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig. 58
: i nti
AN
ny
Uy
C.(Culiciomyia) spathifurca
Bram: Genus Culex in Thailand 167
Integument spiculose or glabrous; hairs 1,2,3-P single, pectinate, subequal
in length and width; 4-6, 8-P well developed. Abdomen. Comb variable,
with from 4 to approximately 50 scales; siphon variable in size and shape,
with from 3 to 7 subventral tufts; 1 or more subventral tufts may or may not
be inserted laterally out of line; pecten restricted to the basal third or less
of the siphon; ventral brush consisting of 10 or more tufts inserted on the
grid; anal gills variable in size and shape; saddle asd aR ringing segment
X except in veolitoralis.
PUPA. Without striking subgeneric chivashan labios,
DISTRIBUTION. Members of the subgenus Culex are distributed
throughout all zoogeographical regions of the world and are the most common
members of the genus Culex. The subgenus is widely distributed throughout
Thailand, but some species are limited by ecological requirements. Ap-
parently no species of the subgenus is endemic to Thailand.
TAXONOMIC DISCUSSION. Females of the subgenus Culex, although
usually exhibiting banding and other features of ornamentation, are frequently
difficult to determine accurately. Many species which show unique features
in the immature stages have extremely similar females, and in these cases
final determinations must depend upon study of the associated larval skins.
The male terminalia of most species are quite distinct, but in certain groups
(e.g. the vishnui subgroup) subtle differences separate species with broadly
different larvae. The fourth stage larva seems to consistently demonstrate
distinctive anatomical characteristics; however, even in this stage, consider-
able variation exists in Some groups, and at times forms with widely
divergent adults have very similar larvae. A case in point is found between
tritaeniorhynchus and mimulus. As adults these species can be separated
by cursory examination owing to the wing scale patterns in mimulus; but as
larvae, the species are extremely similar.
Intraspecific variation in the subgenus is quite extensive wherever
sufficient material is available to permit critical evaluation. C. bitaeniorhynchus
is an example frequently cited as possibly representing a species complex,
due to its extensive: variation in both the adult and immature stages. Evalua-
tion of adult and larval features, however, reveals that this is probably a
plastic species with considerable variation even among siblings.
BIOLOGY. Larvae of Thai species of the subgenus Culex are found
in a wide variety of temporary or semipermanent ground water habitats. Com-
mon habitats include rice paddies, Seepage pools, pot holes, and ditches.
C. pipiens quinquefasciatus is normally found in highly polluted temporary
waters, particularly in urban situations. C. sitiens and veolitoralis are
restricted to brackish water habitats in coastal areas and bitaentorhynchus is
found in association with dense green algal mats.
Biology of adults is even more sketchy than that of the larvae. A
number of species are known to be anthropophilous but host preferences of
most species are poorly understood. Culex is the most important of the sub-
genera from the viewpoint of disease transmission. Two of its members,
tritaeniorhynchus and gelidus, are important vectors of Japanese B encephal-
itis virus in Southeast Asia. Rudnick and Hammon (1961) reported isolating
Figure 58. C. (Culiciomyia) spathifurca. A,B, dorsoventral aspects of
tlie the pupa; C, dorsal aspect of the male terminalia.
168 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C. (Culiciomyia) spathifurca
Bram: Genus Culex in Thailand 169
dengue virus from tritaeniorhynchus in Thailand, but no epidemiological
significance was attached to this report (Rudnick 1966). Chichungunya virus
was isolated from pipiens quinquefasciatus in Bangkok (Rudnick and Hammon
1961), but later work indicated that this observation was not significant
epidemiologically. The same is true of the reported isolation of eastern
equine encephalitis virus from pipiens quinquefasciatus in Bangkok (Rudnick
and Hammon 1961). However, pipiens quinquefasciatus is the most impor-
tant vector of urban filariasis caused by periodic Wuchereria bancrofti
wherever the disease occurs.
Bennett, Warren, and Cheong (1966) and Bennett and Warren (1966)
reported that in Malaya, 2 species of the subgenus Culex, annulus and sitiens,
are natural vectors of Plasmodium juxtanucleare (a parasite of gallinaceous
birds). | Other species of the subgenus, tviiaeniorhynchus, gelidus, and
pseudovishnui, are at least potential vectors, but the parasite failed to
develop in pipiens quinquefasciatus.
KEY TO SPECIES OF THE SUBGENUS CULEX IN THAILAND -
ADULT FEMALES
1. Proboscis without distinct white banding; 1 or 2 lower mesepimeral
bristles present; tarsomeres unbanded....... (pipiens group) 2
Proboscis ringed with a distinct white band; lower mesepimeral
bristles absent; tarsomeres with narrow, pale basal bands......
. (sitiens group) 4
2(1). . Abdomival terga unbanded. . 0... ee Ps fuscocephala (p. 184)
Abdominal terga with narrow, white, basal bands............. 3
3(2). Integument of the pleuron with blackish-brown areas.............
hutchinsoni (p. 188)
Integument of the pleuron usually uniformly pale................
pipiens quinquefasciatus (p. 192)
4(1). Abdominal terga with white apical bands, with apical, lateral
triangular patches and basal bands, or with the distal segments
completely covered with pale scales. ...........-.cc2e200%
(bitaeniorhynchus subgroup) 5
Abdominal terga with white basal bands or, rarely, unbanded;
apical bands or patches not present............ gage Ca ge ate 7
5(4). Wings with pale scales scattered among the dark, particularly on
the costa and subcosta; scutum without silver scales...........
bitaeniorhynchus (p. 258)
Wings without pale scales scattered among the dark; scutum with
Figure 59. C. (Culiciomyia) spathifurca. Fourth stage larva: dorso-
é ventral view of the head, thorax and abdomen, and lateral
aspect of the terminal abdominal segments.
170 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
L fe
Bip
Bram: Genus Culex in Thailand 171
silver scales anterior to the prescutellar space............. 6
6(5). Apical pale tergal bands with the proximal margin straight; basal
ter@al bands very narrow..:6.0.5) 6 er ee is sinensis (p. 268)
Apical pale tergal bands represented on the proximal segments
by triangular lateral patches which may converge at the
center; basal tergal bands rather broad. . .pseudosinensis (p. 265)
7(4). Scutum densely covered with distinct, silvery scales, at least
anterior to the prescutellar space......... (selidus subgroup) 8
Scutum uniformly brown or with some golden or yellow scales..... 9
8(7). Silver scaling on the scutum terminating at the level of the wing base;
basal abdominal bands reaching to the lateral edges of the terga,
frequently with a v-shaped median, posterior projection.........
gelidus (p. 24'7)
Silver scaling on the scutum continued posteriorly through the
prescutellar space and onto the middle of the scutellum; basal
abdominal bands not reaching the lateral edges of the
(OYA SS ee OR ae ee Oe ae ee whitmorei (p. 253)
9(7). Wings with 2 or more distinct patches of light scales on the costa
and subcosta, and with some variable patches on the other
VOU es ee ee a Oe a a Ge mimulus (p. hate
10(9). Anterior surface of the mid and hind femora and tibiae with
distinct, pale s@ipes bo Oe Vee es barraudi (p. 209)
Mid and hind femora and tibiae without pale stripes on the
DOUErIOR SUriace ee nl oe Oe ne ee 11
11(10). Proboscis usually with accessory pale patches proximal to the
median pale band on the ventral surface; teeth of the bucco-
pharyngeal armature very long and filamentous; erect scales
on the occiput dark brown; usually a small, dark brown
Cee ee ee a ee en tritaeniorhynchus (p. 2'75)
Proboscis without accessory pale patches on the ventral surface;
teeth of the buccopharyngeal armature short, not filamentous;
erect scales on the occiput varying from pale golden to dark
brown; usually larger species 2 ee ee eo Wa eo es 12
12(11). Erect scales of the vertex uniformly dark brown, or dark brown
on the occiput and with several almost black scales postero-
BY in cece ais we al Eg ecu Wig tan aoe yeaa wy ame a 13
Figure 60. C. (Culiciomyia) spiculothorax. Fourth stage larva: dorso-
ventral view of the head, thorax and abdomen, and lateral
— aspect of the terminal abdominal segments.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
172
serena
a
¥
‘a) termi
¥
iciomyta
C.(Culi
Bram: Genus Culex in Thailand 173
Erect scales of the vertex yellowish to golden brown on the occiput,
dark brown to black posterolaterally................0-.28- 14
13(12). Anterior surface of the mid femur speckled with brown and white
scales; mid and hind tibiae dark with a trace of pale scales
forming somewhat of astripe............... sitiens (p. 239)
Anterior surface of the mid femur predominantly dark, slightly
lighter on the ventral margin, but without white speckling;
mid and hind tibiae not as above....... annulus (in part, p. 205)
14(12). Abdominal terga II-VI without basal pale bands................
perplexus (in part, p. 221)
Abdominal terga II-VI with basal pale bands... . neolitoralis (p. 236)
pseudovishnui (p. 222)1
whitei (p. 233)
alienus (p. 202)
perblexus (in part, p. 221)
annulus (in part, p. 205)1
KEY TO SPECIES OF THE SUBGENUS CULEX IN THAILAND -
ADULT MALES
‘. Arms of the outer division of the phallosome simple, nearly
parallel in dorsal aspect; arms of the inner division simple,
curved laterally; basal sternal process of the proctiger very
short, straight, usually lightly pigmented..................
pipiens quinquefasciatus (p. 192)
Arms of the outer division of the phallosome widely divergent;
arms of the inner division usually with numerous teeth (except
in hutchinsoni, fuscocephala, and sinensis); basal sternal
process of the proctiger long and curved, usually strongly
pigmented (except in neolilovalis) oO... is Vi eK WE US we bs 2
2(1). Subapical lobe of the basimere distinctly divided, the 3 basal rods on
a tubercle separated from the leaf and accessory setae; arms
1 Of the 6 species which cannot be conclusively recognized on the basis of the
adult female, pseudovishnui and annulus are undoubtedly the most common in
Thailand according to larval collections.
Figure 61. C. (Culiciomyia) termi. Adult habitus: A, lateral aspect
of the male head; B, lateral aspect of the female head and
en thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female
abdomen.
WO 2, nocl, 1967
Contrib. Amer. Ent. Inst.
174
C. (Culiciomyia ) termi
Bram: Genus Culex in Thailand 175
of the inner division of the phallosome simple, but with faint
striations or minute denticles, and truncate distally...........
fuscocephala (p. 184)
Subapical lobe of the basimere undivided; arms of the inner
division of the phallosome usually complex, if simple without
Leabnt, SUr Tae isl OT ak Oe ee PRG 3
3(2). Distimere expanded medially; inner division of the phallosome
short, slightly curved laterally, ahd heavily sculptured; outer
division shorter, broad, bluntly rounded and spiculose.........
gelidus (p. 247)
Distimere not noticeably expanded medially; inner and outer
divisions of the phallosome not aS above. ..........c2-2ece00- 4
4(3). Inner division of the phallosome simple, without denticles........ 5
Inner division of the phallosome complex with at least some
denticles; when observed in lateral aspect, with a spiculate
portion (except in whitmorei and whitei).........00+200. ues
5(4). Arms of the inner division of the phallosome straight, parallel,
minutely spiculose; outer division greatly reduced, represented
by a small basal protuberante . 04:0. 26 Vea sinensis (p. 268)
Arms of the inner division of the phallosome simple, prominently
curved and directed laterally; outer division complex, with
some laterally directed processes........ hutchinsoni (p. 188)
6(4). Basal rod of the subapical lobe of the basimere considerably longer
and broader than the other 2 rods; inner division of the phallo-
some without a spiculate portion........... whitmorei (p. 253)
Basal rod of the subapical lobe of the basimere shorter than the
other 2 rods, if longer, only slightly so; inner division of the
phallosome with a spiculate portion (except in whitei)......... 7
7(6). Inner division of the phallosome with from 2 to 4 large, bluntly
rounded denticles, the spiculate portion very prominent with an
extended projection directed away from the denticles; leaf-like
seta on_the subapical lobe of the basimere short............. 8
Inner division of the phallosome with 3 or more smaller, sharply
pointed denticles; the spiculate portion without an extended
projection directed away from the teeth (except in neolitoralis
and sitiens); leaf-like seta on the subapical lobe of the basimere
moderate to large in breadth and length.................. 9
8(7). Basal sternal process of the proctiger represented by a bluntly
rounded protuberance; leaf on the subapical lobe of the basimere
WONT SIONOOR oi os ha oi Ue rae ee bitaeniorhynchus (p. 258)
Figure 62. C. (Culiciomyia) termi. A,B, dorsoventral aspects of the
ecw pupa; C, dorsal aspect of the male terminalia.
176 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Bram: Genus Culex in Thailand itt
Basal sternal process of the proctiger short, but distinctly
developed and gently directed posteriorly; leaf on the subapical
lobe of the basimere somewhat broader. . .pseudosinensis (p. 265)
9(7). Outer division of the phallosome with a series of at least 3 denticles.
similar in size and shape to those of the inner division; subapical
lobe of the basimere with 4 accessory setae next to the leaf......
whitei (p. 233)
Outer division of the phallosome without a series of denticles
similar to those of the inner division; subapical lobe of the
basimere usually with 3 accessory setae next to the leaf...... 10
10(9). Spiculate portion of the inner division of the phallosome with a
distinct heel or projection on the inner margin; all 3 accessory
setae next to the leaf on the subapical lobe of the basimere sub-
equal in Tenet and hooked . o. 60 ks ee ne al 11
Spiculate portion of the inner division of the phallosome smoothly
rounded, without a heel or projection on the inner margin;
accessory Setae next to the leaf on the subapical lobe of the
basimeré N0l 26 a00V Gi cu Ge OO i ke oe 12
11(10). Spiculate portion of the inner division of the phallosome with a
prominent projection exhibiting fine denticles; basal sternal
process of the proctiger rather short and straight............
neolitoralis (p. 236)
Spiculate portion of the inner division of the phallosome with a small
spiculate heel on the inner margin; basal sternal process of the
proctiger lone and curved. 24. 46 6 4h a es sitiens (p. 239)
12(10). Wings with 2 or more distinct patches of light scales on the costa
and subcosta, and with some variable patches on the other
WING WOING ics so be ee ee mimulus (p. 213)
Wings without distinct patches of white scales............... 13
13(12). Anterior surface of the mid and hind femora and tibiae with distinct
lonvitudinal pale stripes. 2... 446 os 6. SO _. barraudi (p. 209)
Mid and hind femora and tibiae without longitudinal pale stripes :
on the:anterior surigecew 35 06 8 ew ee alienus (p. 202)
annulus (p. 205)
perplexus (p. 221)
pseudovishnui (p. 222)
: Colless (1957) has suggested that pseudovishnui does not usually possess the
long ventral hairs at the base of the band on the proboscis which are present
in the other closely related species. This character appears to be consistent,
although in some specimens of pseudovishnui 1 or 2 shorter hairs may be
present.
LLL LC LT CL perenne eengunoespneeneoneneneeeae
Figure 63. C. (Culiciomyia) termi. Fourth stage larva: dorsoventral view
‘allie of the head, thorax and abdomen, and lateral aspect of the
terminal abdominal segments.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
178
S|
o
2a TH
w
=
=
=
i,
tin 4
TA)
iy
¢> “| OS bys
RENT TT
C. (Culiciomyia ) thurmanorum
13(12).
3(1).
4(3).
5(4).
6(3).
Bram: Genus Culex in Thailand 179
CECE LE ONS ES ats ene tritaeniorhynchus (p. 27 5)
KEY TO SPECIES OF THE SUBGENUS CULEX IN THAILAND -
FOURTH STAGE LARVAE
Head hair 1-C fine, tapering and filamentous distally........... 2
Head hair 1-C thicker, acuminate, truncate, or bluntly pointed.... 3
Siphon index approximately 5:1 or greater; head hairs 5, 6-C with
2 or 3 branches. seas as 24 Bike eee fuscocephala (p. 184)
Siphon index approximately 3:1, the siphon expanded medially; head
hairs 5,6¢C with.5.0r more DPANCHESs ais, 6 iiic efi 8) thu Asa aniga & 8
_ pipiens quinquefasciatus (p. 192)
Head hair 1-C lightly pigmented, long, of even width to apex; pecten
inconspicuous, restricted to the basal fifth of the siphon or
TOSS ku ae a ee ea ida Bike ee ee a &
Head hair 1-C strongly pigmented, shorter, tapering to apex or
short and truncate; pecten obvious, extending through approxi-
mately the basal 1/A.or 1/3.of the Siphon «ioc «656 8 98 ny we a 6
Lateral teeth of the mentum robust, distinctly separated and less
than 10 on each side; thoracic hair 4-P short, simple.........
sinensis (p. 268)
Lateral teeth of the mentum numerous, extremely short and
compact; thoracic hair 4-P long, pectinate................ 5)
Siphon usually with 4 pairs of subventral tufts; individual comb
scales elongated; pecten extending only a very short distance
frony the ase fb iidce wins ee SA te bitaeniorhynchus (p. 258)
Siphon with 3 pairs of subventral tufts, the basal tuft frequently
not paired; individual comb scales relatively short; length of
the pecten somewhat greater than above. . pseudosinensis (p. 265)
Head hair 1-C broad and somewhat flattened, its apex rounded or
irregular; anal gills bulbous, not as long as the saddle........ 7
Head hair 1-C slender or moderately thickened, its apex acuminate;
: In tritaeniorhynchus, the spiculate portion of the inner division of the
phallosome usually exhibits a slightly expanded apex which projects some-
what beyond the attachment of the teeth (see insert on figure 77C); however,
if the specimen is not positioned exactly, this feature may be distorted, and
must therefore be used with caution.
Figure 64. C. (Culiciomyia) thurmanorum. A,B, dorsoventral aspects
<j
of the pupa; C, dorsal aspect of the male terminalia.
180 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C. (Culiciomyia ) thurmanorum
7(6).
8(6).
9(8).
10(9).
11(10).
42(11).
13(8).
14(13).
Bram: Genus Culex in Thailand 181
Anal saddle completely ringing segment X; pecten tooth with ap-
proximately 9 fine, lateral barbs............ sitiens (p. 239)
Anal saddle incomplete, consisting of separate dorsal and ventral
plates; pecten tooth with from 2 to 7 coarse lateral barbs.......
neolitoralis (p. 236)
Individual comb scales fan-shaped, fringed with subequal spicules,
or rarely with the median distal spicule only slightly longer
than the dateral spicules. 5.50) 2G ee ae 9
At least some of the individual comb scales with the median distal
spine obviously longer and broader than the lateral spicules... .13
Siphon with a single, prominent, subapical spine...............
hutchinsoni (p. 188)
Siphon without a prominent, subapical spine................ 10
Siphon greatly expanded medially; subventral tufts of the siphon
all inserted ina straight line.............. gelidus (p. 247)
Siphon not greatly expanded medially; 1 or more pairs of subventral
tufts of the siphon inserted laterally out of line............. 11
Thoracic hair 4-P single; median distal spicule of the comb scale
occasionally slightly longer than the lateral spicules..........
mimulus (p. 213)
Thoracic hair 4-P bifid; median distal spicule of the comb scale
never longer than the lateral spicules................0.2.- 12
Stirrup-shaped piece of the siphon without a proximally directed
prolongation; abdominal hair 1-X with 3 or 4 short branches.....
tritaeniorhynchus (p. 2'75)
Stirrup-shaped piece of the siphon with a prolongation of the
ventral cornu directed proximally; abdominal hair 1-X usually
with 2 fairly long, fine branches........... barraudi (p. 209)
Subventral tufts of the siphon bifid, pectinate, with 2 additional
shorter, simple pairs inserted laterally out of line...........
whitmorei (p. 253)
Subventral tufts of the siphon with 2 or more branches which are
always simple, usually with only 1 pair inserted laterally
out eof Aine. ie he ee a ee I a ee a, 14
Subventral tufts of the siphon usually with 6 or more branches;
Figure 65. C. (Culiciomyia) thurmanorum. Fourth stage larva: dorso-
ae
ventral view of the head, thorax and abdomen, and lateral
aspect of the terminal abdominal segments.
182 Contr ib. baie Ent. Inst., vol. 2, no. 1, 1967
thoracic integument minutely spiculose........ annulus (p. 205)
Subventral tufts of the siphon usually with 5 or less branches;
HOF ACC TRLESUINGN! CIADTONS 0 ki We ie eee ek be ke 19
15(14). Comb consisting of approximately 40 scales; subventral tufts equal
to or less than the width of the siphon at the point of insertion,
peuaily abide 2k Die CW A OS. whitei (p. 233)
Comb consisting of 25 or less scales; length of the subventral tufts
greater than the width of the siphon at the point of insertion, at
least some of the tufts with 3 or more branches............ 16
16(15). Head hairs 5,6-C usually bifid; the length of the basal subventral
tuft greater than 3 times the width of the siphon at the point of
TRB OCUR CR ei eal) Me a ies ha ee ti alienus (p. 202)
Head hair 5-C with from 3 to 5 branches, 6-C with 2 or 3
branches; length of the basal subventral tuft less than 2 times
the width of the siphon at the point of insertion............. 17
17(16). Comb consisting of from 5 to 13 long, sharp scales arranged in an
irregular row; head hair 4-C usually bifid or trifid; abdominal
hairs 6-V, VI with 3 branches; head hair 1-C never with
accessory lateral spicules............ pseudovishnui (p. 222)
Comb consisting of from 12 to 20 rather short scales arranged
in 2 or 3 irregular rows; head hair 4-C usually single; ab-
dominal hairs 6-V, VI bifid; head hair 1-C frequently with
accessory lateral spicules. ............06- perplexus (p. 221)
PIPIENS GROUP
ADULT. Head. Proboscis without a median band of pale scales.
Thorax. At least 1 lower mesepimeral bristle present; legs without pale
bands on the tarsomeres. Abdomen. Terga with or without pale basal bands.
LARVA. Head, Hair 1-C usually fine and filamentous, except in
hutchinsoni which exhibits a prominent subapical spine on the siphon.
DISTRIBUTION. Species of this group are distributed primarily in
the Ethiopian region and the New World, with only a few representatives
in the Oriental region. The species found in Thailand are usually common
and widely distributed, except for hutchinsoni which apparently has a more
limited distribution.
TAXONOMIC DISCUSSION. Edwards (1932) included fatigans (= pipiens
quinquefasciatus) in the pipiens series of group B; fuscocephala and
hutchinsoni were included with species of uncertain position in group B. The
pipiens group, as here recognized, encompasses Edwards' group B without
Figure 66. C. (Culex) fuscocephala. Adult habitus: A, lateral aspect
of the male head; B, lateral aspect of the female head and
thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
—>
C. (Culex ) fuscocephala
Bram: Genus Culex in Thailand
184 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
recognizing subgroups. Of the 3 species found in Thailand, pipiens
quinquefasciatus and fuscocephala fall within the pipiens series of Edwards
due to the presence of 1 or more lower mesepimeral bristles in the adult
female and the absence of a median pale band on the proboscis. C.
hutchinsoni possesses the distinctive characters of the adult, but head hair
1-C in the fourth stage larva is robust and acuminate in contrast to the
filamentous form found in the other 2 species.
CULEX (CULEX) FUSCOCEPHALA THEOBALD 1907
(Figures 66, 67, and 68)
Culex fuscocephala Theobald 1907, Mon. Cul. 4: 420 (¢); Theobald 1910,
Mon. Cul. 5: 363 (distribution); Barraud 1924, Indian J. med. Res.
11: 1270. (¢*, 9).
Culex uniformis Leicester 1908, Cul. Malaya: 159 (¢); Edwards 1917, Bull.
ent. Res. 7: 225 (synonymy).
Culex minimus Leicester 1908, Cul. Malaya: 160 (9, “); Edwards 1932, in
Wytsman, Genera Insect. fasc. 194: 212 (synonymy).
Culex taytayensis Banks 1909, Philipp. J. Sci. 4: 545 (o, %); Edwards 1913,
Bull. ent. Res. 4: 234 (synonymy).
Culex luteola Theobald 1910, Mon. Cul. 5: 378 (@*); Edwards 1913, Bull.
ent. Res. 4: 234 (synonymy).
Culex inelegans Dyar 1920, Insec. Inscit. menst. 8: 179 (*); Barraud 1934,
Fauna Brit. India, Diptera 5: 424 (synonymy).
Culex fuscitarsis Barraud 1924, Indian J. med. Res. 11: 1272 ('*, 9).
NEW SYNONYMY.
Culex (Culex) fuscocephalus Theobald: Barraud 1934, Fauna Brit. India,
Diptera 5: 424 (o*, 9, L*); Bonne-Wepster and Brug 1937, Geneesk.
Tijdschr. Ned. -Ind. 77: 80 (“, 2?*); Bonne-Wepster and Brug 1939,
Geneesk. Tijdschr. Ned. -Ind. 79: 1277 (L*); Bohart 1945, Navmed
580: 78 («*, L); Bonne-Wepster 1954, Roy. trop. Inst. Amst. Spec.
Pub. 111: 133 (o, 9*, L*); Lien 1962, Pacif. Ins. 4: 633 (distribu-
tion); Safyanova et al. 1964, Zool. Zhur. 43: 1177 (distribution).
Culex (Culex) fuscocephala Theobald: Delfinado 1966, Mem. Amer. ent.
Inst. 7: 143 (o*,¢?, L).
The adult female may be recognized by the unbanded abdominal terga
and by the presence of dark areas on the integument of the pleuron. In the
adult male, the distinctive phallosome is diagnostic. The fourth stage larva
is distinguished by chaetotaxy of the head and by the characteristic siphon.
FEMALE. A moderate to small sized, exceptionally dull species.
Head. (Figure 66B). Proboscis bronze-brown, without light bands or
scattered light scales; palpus similar in color to the proboscis; decumbent
scales of the vertex white; erect scales dark brown. Thorax. (Figure 66B,
E). Scutum covered with bronze-brown scales, but with somewhat lighter
scales in the area of the anterior pronotal lobe and the prescutellar space;
Figure 67. C. (Culex) fuscocephala. A,B, dorsoventral aspects of the
pupa; C, dorsal aspect of the male terminalia. ;
or
Jy) Me |
Thy vey ad a LE
! Me 8s
C.(Culex) fuscocephala
a a
2” Tf
eal)
ze | oj
TL) Ww meth A
(etre hae
Bram:
Genus Culex in Thailand
186 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
integument of the pleuron basically light brown, but with variable dark patches
on the mesepimeron, sternopleuron, and posterior spiracular plate; 1 or 2
lower mesepimeral bristles present; patches of light scales on the upper mesep-
imeron and upper and posterior sternopleuron. Wing. (Figure 66C). Legs.
(Figure 66D). Hind femur virtually all white on the anterior surface, but with
a very dark band at the apex which extends proximally along the dorsal margin;
hind tibia and tarsus uniformly covered with bronze-brown scales; fore and
mid legs bronze-brown, but with varying amounts of light scales on the
anterior surface of the femora. Abdomen. (Figure 66F). Terga completely
bronze-brown, but with some faint suggestion of narrow light basal bands on
the proximal terga of some specimens; sterna light brown.
: MALE. Head. (Figure 66A). Palpus uniformly dark, but in some
specimens somewhat lighter on the distal segments. Terminalia. (Figure
67C). Subapical lobe of the basimere well developed, distinctly divided, the
3 basal rods on a tubercle separated from the leaf and with a rather strong,
bluntly rounded seta set on another small independent tubercle; distimere
characteristically curved, with minute annulations on the convex surface of
the apex; outer division of the phallosome consisting of 4 or more strong
denticles, inner division rather long and slender, covered with faint stria-
tions or minute denticles; proctiger crowned with a strong tuft of spines,
with 3 cercal setae present; basal sternal process strong and curved.
PUPA. (Figure 67A,B).
LARVA. (Figure 68). Head. Antenna with a narrow, dark basal
ring and often somewhat darker beyond insertion of hair 1-A; head hair 1-C
filamentous, its length greater than half the distance between the bases of
the pair; 4-C single, simple, very fine and filamentous; 5,6-C usually bifid
or trifid, pectinate. Thorax. Integument glabrous; hairs 1,2, 3-P single,
pectinate, subequal in length; 4-P usually bifid, occasionally single, pecti-
nate; 5,6-P single, pectinate; 7,8-P with 2 or 3 branches, pectinate; 14-P
single, simple. Abdomen. Integument glabrous; comb consisting of from 35
to 40 fan-shaped scales arranged in a triangular patch; siphon index variable,
ranging from 4. 8:1 to 7:1; 3 or 4 pairs of subventral tufts inserted on the
siphon beyond the pecten, the subapical tuft inserted laterally out of line; in-
dividual tufts with from 2 to 4 branches, their length less than the width of
the siphon at the point of insertion; pecten consisting of from 10 to 12 teeth
restricted to the basal third of the siphon; individual pecten tooth with a
strong, narrow, pointed distal spine and from 1 to 3 strong lateral barbs;
saddle completely ringing segment X.
TYPE DATA. Holotype female of fuscocephala from Peradeniya,
Ceylon in the British Museum. The type locality of uniformis is Batu Gajah,
(Perak), Malaya, but the type specimen is non-existent. The type locality of
minimus is Kuala Lumpur, (Selangor), Malaya, but the type specimen is non-
existent. The holotype of tayiayensis has been destroyed, but a paratype male
(terminalia mounted and attached to pin) from Taytay, Philippine Islands is in
the British Museum. Holotype female of luteola from Peradeniya, Ceylon in
the British Museum. Holotype male of inelegans (terminalia slide- mounted)
Figure 68. C. (Culex) fuscocephala. Fourth stage larva: dorsoventral
view of the head, thorax and abdomen, and lateral aspect of
the terminal abdominal segments. —e
187
Bram: Genus Culex in Thailand
Fig.68
Z
3
1 8 A
12 a
: 119 Z
‘ , :
VB |i , | 13 [ [ y f
ve ae
Ae a4
C. (Culex) fuscocephala
188 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
from Los Banos, Philippines in the U. S. National Museum. Lectotype male
of fuscitarsis hereby designated: syntype male (terminalia slide-mounted),
'no. 1701, Pachmari, India, July 1915, V. H. Dowson, B.M. 1923-585", in
the British Museum.
DISTRIBUTION. In THAILAND, this species is common and probably
widely distributed throughout the country and specimens have been examined
from: Ayutthaya, Chiang Mai, Chiang Rai, Chon Buri, Krung Thep, Lampang,
Lamphun, Mae Hong Son, Nakhon Nayok, Nakhon Ratchasima, Narathiwat,
Pathum Thani, Phitsanulok, Phare, Sara Buri, Surat Thani, Thon Buyvi,
Ubon Ratchathani, and Udon Thani. The species has also been reported from
CEYLON, INDIA, NEPAL, BURMA, CHINA, TAIWAN, PHILIPPINES,
INDONESIA, MALAYA, ANDAMAN ISLANDS, and INDOCHINA. Specimens
have been examined in the British Museum collection from Kota Belud,
NORTH BORNEO**,
During this study the following specimens from Thailand have been
examined: 1,044 females and 395 males, 9 of which were individual rear-
ings, and 133 larvae.
TAXONOMIC DISCUSSION. In the original description of fusciiarsis,
Barraud (1924a) indicated that the species differed slightly from fuscocephala
in the form of the male terminalia and by possessing pale narrow, ochreous
bands on the abdominal terga. Examination of an extensive series of
fuscocephala has revealed that these differences cited by Barraud fall well
within the normal range of variation of fuscocephala. Comparison of the
type specimens of both species confirms the conclusion that fuscitarsis and
fuscocephala are conspecific.
BIOLOGY. Larvae are found primarily in various ground water
habitats such as rice fields, ground pools of various sizes, impounded water,
and other semipermanent aquatic habitats usually with emergent vegetation.
The adults of both sexes are readily attracted to light, and adult females have
been reported feeding on man and buffalo. Hu (1958) reported a positive iso-
lation of Japanese B encephalitis from this species in Taiwan.
CULEX (CULEX) HUTCHINSONI BARRAUD 1924
(Figures 69, 70, and 71)
Culex hutchinsoni Barraud 1924, Indian J. med. Res. 11: 1261 (o*, 9). _
Culex (Culex) hutchinsoni Barraud: Barraud 1934, Fauna Brit. India,
Diptera 5: 423 (o'*, 2); Colless 1955, Ann. trop. Med. Parasit. 49:
316 (¢, 2, L*); Thurman 1959, Univ. Md. Agric. Exp. Sta. Bull.
A-100: 122 (distribution).
The adult female may be recognized by the markings on the integument
of the pleuron and by the abdominal banding patterns. The distinctive phallo-
Figure 69. C. (Culex) hutchinsoni. Adult habitus: A, lateral aspect of
the male head; B, lateral aspect of the female head and
thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
=
189
Genus Culex in Thailand
Bram
insont
e
C. (Culex) hutch
190 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
some of the male terminalia is diagnostic. In the fourth stage larva, the
Single subapical spine on the siphon clearly distinguishes this species from
all other Oriental members of the genus.
FEMALE. Head. (Figure 69B). Proboscis without distinct light
bands or scattered light scales but with the ventral scales and those of the
proximal half somewhat paler than the distal scales; palpus dark brown,
usually without pale scales; decumbent scales of the vertex light brown,
becoming lighter at the orbital line; erect scales dark brown. Thorax.
(Figure 69B,E). Scutum sparsely covered with bronze-brown scales and
with indistinct but slightly darker areas along the acrostichal line and in the
area of the fossa; integument of the pleuron light brown, with indistinct and
variable dark bands across the upper and lower sternopleuron and mesepimeron;
small patches of light scales on the upper and posterior sternopleuron and
middle mesepimeron; 1 lower meSepimeral bristle present. Wing. (Figure
69C). Legs. (Figure 69D). Anterior surface of the hind femur with a white
longitudinal stripe on the ventral half and dark dorsally and apically; hind
tibia dark with a narrow apical white band and somewhat lighter on the ventral
margin; hind tarsus completely dark; fore and mid legs similar in color to
the hind legs. Abdomen. (Figure 69F). Terga dark scaled with rather nar-
row, variable and sometimes indistinct, pale basal bands on segments II-VU;
tergum VIII usually completely pale; sterna uniformly covered with pale
golden-brown scales.
MALE. Head. (Figure 69A). Proboscis with a narrow median band
of pale scales; palpus dark brown, with a rather broad, but variable subapical
pale band on segment II]; frequently with variable amounts of basal pale
scaling on segments IV and V. Terminalia. (Figure 70C). Subapical lobe of
the basimere well developed, with 3 strong basal rods followed by 3 hooked
accessory setae, a broad, striated leaf and a gently curved seta; distimere
normal in shape, without distal annulations; outer division of the phallosome
very broad, and tapering gradually to a point, but without noticeable denticles
and with the basal arm well developed and tapering to a sharp point; inner
division narrow, acutely angled and tapering to a sharp point; proctiger
crowned with a strong tuft of spines, with 2 or 3 cercal setae present; basal
sternal process strong and curved.
PUPA. (Figure 70A,B).
LARVA. (Figure 71). Head. Antenna with a very narrow dark basal
ring; hair 1-A inserted somewhat beyond the middle of the shaft; head hair
1-C darkly pigmented, robust, tapering gradually to a sharp point, its length
approximately half the distance between the bases of the pair; 4-C single,
Simple; 5,6-C usually trifid, pectinate. Thovax. Integument covered with a
random pattern of very minute spicules (not illustrated); hairs 1,2,3-P single,
pectinate, 3-P considerably shorter than 1,2-P; 4-P with 4 or 5 pectinate
branches; 5,6-P single, pectinate; 7, 8-P bifid, pectinate; 14-P single, simple.
Abdomen. Integument glabrous; comb variable, consisting of approximately
35 fan-shaped scales arranged in 3 irregular rows; siphon index variable,
ranging from 2.5:1 to 3.5:1, enlarged basally and tapering rapidly at the
distal third; a single, very prominent subapical spine projecting from the
Figure 70. C. (Culex) hutchinsoni. A,B, dorsoventral aspects of the
pupa; C, dorsal aspect of the male terminalia. .
Bram: Genus Culex in Thailand 191
. i)
\ i}
1 7 Ls
\\ 4 ee: ) hy
vy Wash Uy PO yA
k ag) PTT
~
C. (Culex) hutchinsoni
192 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
lateral margin of the siphon; 4 subventral tufts inserted on the siphon, the
subapical tuft laterally out of line; individual tufts with from 2 to 6 branches,
their length less than the width of the siphon at the point of insertion; pecten
consisting of from 4 to 8 teeth restricted to approximately the basal third to
half of the siphon; individual pecten tooth elongated, with a long, narrow,
sharply pointed apical spine and from 1 to 3 short, strong basolateral barbs;
saddle completely ringing segment X.
TYPE DATA. Lectotype of hutchinsoni hereby designated: syntype
male, "India, Nongpoh, Assam, VII. 1922, P. J. Barraud, B.M. 1923-585,
1726", in the British Museum.
DISTRIBUTION. In THAILAND, specimens of hutchinsoni have been
collected from: Chiang Mai, Lampang, Nakhon Ratchasima, Nan, and Thon
Buri. This species has also been reported from Assam, INDIA, Selangor,
MALAYA, and SINGAPORE. Specimens have been received in the U. S.
National Museum from An Khe, SOUTH VIETNAM**, and the author has seen
material in the British Museum from Nampam, Shan States, BURMA**,
During this study the following material was examined: 22 females
and 19 males, 12 with their associated larval and pupal skins, and 78 larvae.
TAXONOMIC DISCUSSION. The adult female may possibly be confused
with pipiens quinquefasciatus, but markings on the integument of the pleuron
and the lighter scaling of the proboscis are usually diagnostic. In the adult
male, the distinctive phallosome clearly separates this species from all other
members of the subgenus. The fourth stage larva is immediately recognized
by the single subapical spine on the siphon. Other Oriental species of the
subgenus Culex which exhibit spines on the siphon include mimeticus, jacksont,
and an undescribed species from China, but in each instance more than 1 spine
is present.
BIOLOGY. Colless (1955) reported that breeding habits of hutchinsoni
resemble those of pipiens quinquefasciatus and that adults appear to frequent
houses. Collections from Thailand do not confirm these observations, and
larvae have been collected from pools, rock holes, stream pools, and elephant
track depressions. Larvae were collected in South Vietnam from a ground
pool.
CULEX (CULEX) PIPIENS QUINQUEFASCIATUS SAY 1823
(Figures 72, 73, and 74)
Culex quinquefasciatus Say 1823, J. Acad. nat. Sci. Phila. 3: 10 (adult).
Culex fatigans Wiedemann 1828, Aussereurop. Zweifl. Inseckten 1: 10 (¢, 2);
Stone 1956(1957), Proc. ent. Soc. Wash. 58: 342 (synonymy).
Culex cingulatus Doleschall 1856, Natuurk. Tijdschr. Ned.-Ind. 10: 405
(preoccupied by Fabricius 1805).
Culex doleschallii Giles 1900, Handb. : 338 (new name for cingulatus
Doleschall); Barraud 1934, Fauna Brit. India, Diptera 5: 420
(synonymy).
Figure 71. C. (Culex) hutchinsoni. Fourth stage larva: dorsoventral
view of the head, thorax and abdomen, and lateral aspect of
the terminal abdominal segments. >
193
Genus Culex in Thailand
Bram:
Fig. 71
C. (Culex) hutchinsoni
194 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Culex fouchowensis Theobald 1901, Mon. Cul. 2: 137 (o*, 9*); Barraud
1934, Fauna Brit. India, Diptera 5: 420 (synonymy).
Culex reesi Theobald 1901, Mon. Cul. 2: 145 (o'*, 9*); Barraud 1934, Fauna
Brit. India, Diptera 5: 420 (synonymy).
Culex sericeus Theobald 1901, Mon. Cul. 2: 147 (?*); Barraud 1934, Fauna
Brit. India, Diptera 5: 420 (synonymy).
Culex hensemaeon Dyar 1920, Insec. Inscit. menst. 8: 178 (?); Edwards
1922, Ind. J. med. Res. 10: 278 (synonymy).
Culex (Culex) fatigans Wiedemann: Barraud 1924, Ind. J. med. Res. 11:
1264 (o*, 9); Barraud 1924, Ind. J. med. Res. 12: 430 (L*); Borel
1926, Arch. Insts. Pasteur Indo-Chine 3-4: 24 (o*, 9, L*); Barraud
1934, Fauna Brit. India, Diptera 5: 420 (o*, 9, L*); and some
other British authors.
Culex (Culex) quinquefasciatus Say: Bohart 1945, U. S. Navmed 580: 79
(*, L); LaCasse and Yamaguti 1950, Mosq. Fauna Japan and Korea:
220 (o'*, 9*, L*); Belkin 1962, Mosq. S. Pacif.: 195 (“*, 9, P*, L*);
and some other American authors.
Culex (Culex) pipiens quinquefasciatus Say: Stone, Knight, and Starcke 1959,
Synoptic Cat. Mosq. Wld.: 254 (systematics); and some other
American authors.
NOTE. The above synonymy is incomplete and lists pertinent references for
the Oriental region only; for complete synonymy in all geographical
regions consult Stone, Knight, and Starcke 1959, Synoptic Cat. Mosq.
Wld.: 254, and Stone 1963, Proc. ent. Soc. Wash. 65: 135.
The adult female may be recognized by the absence of a median pale
band on the proboscis, by the banded abdominal terga, and by the pale in-
tegument of the pleuron. The simple phallosome of the male terminalia and
the short basal sternal process of the proctiger are diagnostic. The fourth
stage larva is identified by the chaetotaxy of the head and siphon.
FEMALE. Head, (Figure 72B). Proboscis and palpus dark brown,
without light bands or scattered pale scales; decumbent scales of the vertex
pale; erect scales usually dark, but with some pale scales at the occiput.
Thorax. (Figure 72B,E). Scutum covered with variable pale brown scales,
frequently lighter on the front, sides and on the prescutellar area; integument
of the pleuron pale brown, usually without darkened areas; usually 1, but
sometimes 2 lower mesSepimeral bristles present; small patches of dull white
scales present on the upper meSepimeron, anterior mesepimeron, upper
sternopleuron, and posterior sternopleuron. Wing. (Figure 72C). Legs.
(Figure 72D). Anterior surface of the hind femur with pale scales on the
ventral half, completely covered with pale scales at the base, and completely
dark at the apex; hind tibia dark scaled, with a few pale scales at the apex;
hind tarsus dark; fore and mid legs similar to the hind legs. Abdomen.
(Figure 72F). Terga dark with moderately broad convex white basal bands
Figure 72. C. (Culex) pipiens quinquefasciatus. Adult habitus: A, lateral
aspect of the male head; B, lateral aspect of the female head
and thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
oS
Bram: Genus Culex in Thailand 195
C. (Culex) pipiens quinquefasciatus
196 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
and usually with light basolateral white patches on terga II-VII; sterna pre-
dominantly pale, with variable patches of dark scales.
MALE. (Figure 72A). Antenna shorter than the proboscis; palpus
dark, with a somewhat paler area at the base of segment III, with a ventral
line of white scales on segment IV, and with a basal ventral patch of white
scales on segment V. Terminalia. (Figure 73C). Subapical lobe of the basi-
mere well developed, with 3 basal rods followed by 2 fine, hooked accessory
setae, a broad, blunt seta, a broad leaf, and a long curved Seta; distimere
normal in shape, without distal annulations; arms of the inner division of the
phallosome rather broad, directed laterally and tapering to a point, arms of
the outer division straight, tapering to a sharp point, nearly parallel in dorsal
aspect; proctiger crowned with a strong tuft of spines, with 2 or 3 cercal
setae present; basal sternal process weakly developed, short and straight.
PUPA. (Figure 73A, B).
LARVA. (Figure 74). Head. Antenna with a narrow, dark basal
ring; head hair 1-C filamentous, its length approximately equal to half the
distance between the bases of the pair; 4-C usually single, simple; 5-C
usually with 6 pectinate branches; 6-C usually with 5 pectinate branches.
Thorax. Integument glabrous; hairs 1,2,3-P single, pectinate, subequal in
length; 4,7, 8-P bifid, pectinate; 5,6-P single, pectinate; 14-P single, simple.
Abdomen. Integument glabrous; comb variable, consisting of from 30 to 50
fan-shaped scales arranged in a broad, triangular patch; siphon index usually
approximately 3:1, almost always less than 5:1; the siphon rather broad
medially with 4 pairs of subventral tufts, the subapical tuft placed laterally
out of line; individual tufts with from 2 to 9 branches, their length less than
the width of the siphon at the point of insertion; pecten variable, consisting of
from 5 to 12 teeth restricted to the basal third of the siphon; individual pecten
tooth with a strong apical spine and 3 or 4 strong lateral barbs and occasionally
several short basolateral spines; saddle completely ringing segment X.
TYPE DATA. The type locality of quinquefasciatus is ''Mississippi
River, United States" but the type specimen is non-existent. Syntypes of
fatigans from Indonesia in the Naturhistorisches Museum, Vienna, Austria.
The type locality of cingulatus is Ambarawa, Java but the location of the type
is unknown. Lectotype of fouchowensis hereby designated: type male
(terminalia slide-mounted), "84, Foochow, T. Rennie, 7/8/00, common
native quarters in suburbs", in the British Museum. The type locality of
veesi is Hong Kong, China but the type has apparently been lost. The type
locality of seviceus is Hong Kong, China but the location of the type is un-
known. The holotype female of hensemaeon from Los Banos, Philippines is
the U. S. National Museum.
DISTRIBUTION. This species is cosmotropical and is undoubtedly
distributed throughout THAILAND. During this study the following specimens
from Thailand were examined: over 750 males and over 300 females, 26
with their associated larval and pupal skins, and 568 larvae.
TAXONOMIC DISCUSSION. The use of the name quinquefasciatus in
preference to fatigans is based on considerations outlined by Stone (1956(1957)).
Briefly stated, if both names apply to the same taxon (and they have, in fact,
Figure 73. C. (Culex) pipiens quinquefasciatus. A,B, dorsoventral
aspects of the pupa; C, dorsal aspect of the male terminalia.
: xa
Bram: Genus Culex in Thailand 197
C. (Culex) pipiens quinquefasciatus
198 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
been so generally accepted for a number of years), then the name with
priority in accordance with the International Rules of Zoological Nomencla-
ture is the one to be used. The treatment of quinquefasciatus as a sub-
species of pzpiens Linnaeus, reflects the concept summarized by Mattingly
(1965) that pipiens is a single polytypic species. Hybridization and apparent
gene flow between various identifiable populations tend to strengthen the
polytypic concept and anatomical studies have demonstrated intermediate
forms in limited areas where at least p. pipiens and p. quinquefasciatus are
sympatric (see for example, Bekku 1956, Barr 1957, and Barr 1960).
BIOLOGY. The biclogies of various populations of pipiens
quinquefasciatus have been studied to a greater or lesser degree in virtually
every area of its distribution. A great volume of data on the biology of this
subspecies has accumulated, but unfortunately there still remains critical
gaps in our knowledge. It must be recognized that results of investigations
in one geographical area are not necessarily valid when applied to another
population of the same subspecies in a different geographical area. The
following summary of the biology of pipiens quinquefasciatus is drawn pri-
marily from recent literature dealing with the subspecies in Southeast Asia.
No attempt has been made to include all biological references and emphasis
has been placed on results of investigations carried out at the W. H. O.
Filariasis Research Unit, Rangoon (assuming that the population found in
Rangoon is most similar to that of Thailand).
The biological cycle of pipiens quinquefasciatus begins when the gravid
female searches for an oviposition site, which generally consists of virtually
any body of water in close association with human habitation which contains a
high degree of organic pollution. Ikeshoji (1966b) reported that breeding field
waters (é.g. water in which larvae are already developing) are more inducive
to oviposition than non-breeding waters. An ether extract of an attractant
has been isolated (but not yet identified) which elicits a high response from
normal females but which cannot be discriminated for by antennectomized fe-
males. De Meillon, Sebastian, and Khan (1966) demonstrated that gravid
females exhibit positive geotaxis and as a result, prefer to oviposit at ground
level. The gravid females have been found to show peaks of oviposition and
visitation to breeding places (de Meillon, Sebastian, and Khan 1967a). The
first of these peaks is characterized by a sharp rise just after sunset anda ©
gradual fall; this is followed by a second peak which occurs at about sunrise
with dramatic rise and fall.
Eggs are deposited on the surface of the water in rafts which usually
consist of from 200 to 250 eggs (Horsfall 1955). Schriver and Bickley (1964)
suggested that water temperature is the only environmental factor which in-
fluences hatching of pipiens quinquefasciatus eggs, and determined that the
optimum temperature range for embryonic development is 23.8°9C. to 29. 4°C.
at which 70 percent of the eggs hatch in about 30 hours. They also determined
that embryonic development stops at 37.5°9C. and at 15.3°C., and that egg
diapause is non-existent. De Meillon, Sebastian, and Khan (1967b) found that
in Rangoon the incubation period of an egg raft is 27.111+0.57 hours at a
Figure 74. C. (Culex) pipiens quinquefasciatus. Fourth stage larva:
dorsoventral view of the head, thorax and abdomen, and
lateral aspect of the terminal abdominal segments.
Bram: Genus Culex in Thailand 199
Fig.74
Stas
|
C. (Culex) pipiens quinquefasciatus
200 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
temperature of 28.110. 66°C. , with 92 percent hatch. The eggs of a raft
usually all hatch within 3 minutes following the emergence of the first free
larva.
The larvae feed on a wide variety of organic material including
diatoms, algae, bacteria, decaying vegetable matter, flagellates, etc.
(Horsfall 1955). The duration of the larval stage is dependent on a great
number of environmental variables, most of which are incompletely under-
stood. Kurihara (1963) investigated the effects of temperature, amount of
food, size of containers, and density on larval mortality and longevity.
De Meillon, Sebastian, and Khan (1967b) determined that at a temperature of
28.59-0. 84°C. , the duration of the female larval life is 13524. 4 hours and
for male larvae is 118. 412.4 hours. Thus, it can be seen that, all things
being equal, the male larvae pupate approximately 17 hours before the fe-
male larvae pupate. The duration of pupal life was determined by these same
authors to be 32. 9510.75 hours for males and 34. 1610.74 hours for females
(at 28. 5940. 84°C. ).
Adults emerge from their pupal skins in 2 clearly defined peaks, one
at 19.00 hours and the other at 21.00 hours (de Meillon, Sebastian, and
Khan 1967c). There is a mass exodus of adults of both sexes from the breeding
site at about sunset. Between 17.00 and 20.00 hours an exodus peak is reached
and it was concluded that those adults which emerge from the pupa too late to
take part in the sunset exodus must wait until the following night when the next
opportunity for exodus presents itself. The adults then disperse, and Lind-
quist ef al. (1967) reported that in a highly populous area of Rangoon, P
tagged adults were capable of traveling at least 0.56 miles within 36 hours
after release. Fussell (1964) recovered P°“ tagged adults from distances of
up to 3.5 miles from their release point in Hawaii.
Horsfall (1955) summarized mating behavior and concluded that in-
semination effected by 1 mating is sufficient for the life of the female.
Laboratory studies by Umino (1966) using marker genes with p. molestus and
pb. pallens indicated that females can copulate with 2 or more males and their
offspring from the same egg raft may have different fathers.
The adult females feed primarily on mammalian hosts and in all
probability man is one of the preferred hosts whenever available. In the
Kemmendine area of Rangoon, de Meillon and Sebastian (1967a) found that 93
percent of the females caught indoors had fed on mammal and 6.7 percent on
avian blood or avian and mammal blood, as compared with 89.5 percent and
10.5 percent respectively when caught out of doors. This sharply contrasts
the results of Hammon e/ al. (1945) who collected freshly engorged females
away from houses in California and found that 88 percent had fed on birds and
the remainder had fed on horse, cow, or dog. However, Toumanoff (1935)
reported that 100 percent of blooded females collected in dwellings in Indo-
china contained human blood and that 84 percent caught in stables had fed on
man and only 16 percent had fed on buffalo. Normally, feeding commences at
around 19.00 hours, reaching a peak at 02.00 hours and ending at 06.00 hours
(de Meillon and Sebastian 1967b). Kurihara and Sasa (1965) reported that in
Bangkok biting activity was highest at about midnight. The gonotrophic cycle
of the female (the time from blood feed to oviposition) was found by de Meillon
and Sebastian (1967b) to be 3 days both in the laboratory and in the field in
Rangoon; however, Kurihara and Hayashi (1965) suggested that the gonotrophic
cycle of p. pallens is somewhat longer in the laboratory. Following feeding,
b
Bram: Genus Culex in Thailand 201
the females may be found resting both in and out of doors, and de Meillon,
Myo Paing, Sebastian, and Khan (1967) concluded that there is every reason
to believe that the outside biting population, which is very large, is drawn
from the outside resting population just as the inside biting population is
drawn from the inside resting population. De Meillon, Sebastian, and Khan
(1967d) found that there was extensive outdoor feeding by both sexes on
crushed sugar cane when this was available in Rangoon. Numerous additional
recent references to the biology of pipiens quinquefasciatus may be found in
the summary of a Seminar on the ecology, biology, and control of the Culex
pipiens complex, sponsored by the World Health Organization (WHO/Vector
Control/125. 65).
The importance of pipiens quinquefasciatus as a primary vector of
urban filariasis caused by periodic Wuchereria bancrofti is generally ac-
knowledged. Although this disease is rare in Thailand, it is very common in |
neighboring countries such as Burma, and therefore pipiens quinquefasciatus
is at least potentially dangerous. Recent studies of the relationships between
pipiens quinquefasciatus and Wuchereria bancrofti include those of de Meillon,
Hayashi, and Sebastian (1967), de Meillon, Grab, and Sebastian (1967),
de Meillon and Sebastian (1967c), Yamamoto (1964), and Yamamoto and
Hayashi (1965).
Chickungunya virus has been isolated from pools of pipiens
quinquefasciatus in Thailand (Hammon, Rudnick, and Sather 1960; Robin e al.
1963; Halstead ef al. 1963) and Singharaj et al. (1966) suggested the possibility
of mechanical transfer of the virus. However, Shah et al. (1964) and
Ramachandra Rao (1964) have reported that virus survival and transmission
could not be demonstrated in this species. Eastern equine encephalitis virus
has also been isolated from pipiens quinquefasciatus (Hammon et al. 1960),
but the significance of this obServation remains open to question. Hodes
(1946) and Reeves and Hammon (1946) recorded laboratory transmission of
Japanese B encephalitis virus by pipiens quinquefasciatus.
SITIENS GROUP
ADULT. Head. Proboscis with a median band of pale scales. Thorax.
‘Lower mesepimeral bristles absent. Abdomen. Terga usually with narrow,
pale basal bands; apical pale bands or patches also present in the ;
bitaeniorhynchus subgroup.
LARVA. Head. Hair 1-C never fine and filamentous, usually darkly
pigmented, but in the biiaeniorhynchus subgroup, long, cylindrical, and
lightly pigmented.
DISTRIBUTION. Species of the szizens group are centered in the
Oriental region with some representatives extending into the Australian and
Ethiopian regions, but none are known from the New World. Species of this
group are found throughout Thailand and several species (e.g. trilaeniorhynchus,
annulus, gelidus) are extremely common.
TAXONOMIC DISCUSSION. The presence of the pale band of the
proboscis and the absence of a lower mesSepimeral bristle are the most out-
standing features which separate the two principle groups of the subgenus
Culex. The sitiens group corresponds to group A of Edwards (1932) and in
Thailand is composed of 4 subgroups as follows: vishnui subgroup, sitiens
202 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
‘subgroup, gelidus subgroup, and bitaeniorhynchus subgroup.
VISHNUI SUBGROUP
FEMALE. Thorax. Scutum uniformly brown or with yellow or
golden scales, never with a dense, distinct pattern of silver scales. Abdo-
men. Terga with white basal bands (except in some specimens of perplexus),
never with pale apical bands or patches.
MALE. Distinctive features as indicated above for the female.
Terminalia. Phallosome complex, usually with the spiculate portion of the
inner division smoothly rounded.
LARVA. Head. Hair 1-C darkly pigmented, slender or moderately
thickened, its apex acuminate. Abdomen. Anal gills elongate, at least as
long as the saddle, usually longer; saddle completely ringing segment X.
TAXONOMIC DISCUSSION. Edwards (1932) included most of the
Species presently recognized as comprising the vishnui subgroup within his
sitiens-Series, and 1 species, mimulus, was included in the mimeticus-
Series. It is felt that the present reassignment of subgroups more accurately
reflects affinities based on all stages.
Members of this subgroup are extremely difficult to recognize on the
basis of the adult males and females alone. Similarities and overlaping
variation render field identification of adults virtually impossible without -
access to their associated larval skins. For this reason, the necessity for
individual rearings with associated larval skins cannot be overemphasized.
CULEX (CULEX) ALIENUS COLLESS 1957
(Figure 75)
Culex (Culex) alienus Colless 1957, Ann. trop. Med. Parasit. 51: 97 (o,?, L*).
The adults cannot be conclusively distinguished from several other
members of the subgroup at the present time. The fourth stage larva may
be recognized by number and shape of the comb scales, by the 2 to 5 branched
subventral tufts of the siphon, and by head hairs 5, 6-C which are usually :
bifid.
FEMALE. Head, Proboscis dark brown with a median band of pale
scales; palpus somewhat darker than the proboscis, tipped at the apex with
one to several randomly placed pale scales; decumbent scales of the vertex
creamy white; erect scales similar in color to decumbent scales (in a male
paratype the scales of the vertex are not as described above, but are almost
identical to the condition found in pseudovishnui). Thorax. Scutum basically
dark brown, but with a variable indistinct grouping of pale golden and silver
Figure 75. C. (Culex) alienus. A, dorsoventral aspect of the head and
prothorax of the fourth stage larva; B, lateral aspect of the
terminal abdominal segments of the fourth stage larva; C,
dorsal aspect of the basimere, lateral aspect of the phallo-
some of the male terminalia.
he
Bram: Genus Culex in Thailand 203
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C. (Culex) alienus
204 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
scales which are particularly noticeable along the lateral margins, the
dorsocentral areas, the supraalar area, and the prescutellar area; pale
scales extend onto the midlobe of the scutellum. Wing. All dorsal wing
scales uniformly light brown. Legs. Anterior surface of the hind femur
predominantly pale with a broad, subapical dark band which extends proximally
along the dorsal border to form a stripe and with a small apical patch of pale
scales; hind tibia dark with a pale apical band; hind tarsus dark, with narrow
pale basal bands on all tarsomere; fore and mid femora dark with a small
apical tuft of pale scales; fore and mid tibiae and tarsi marked as in the hind
legs. Abdomen. Terga dark, with variable but usually moderately broad
convex pale basal bands on segments II-VII and a median basal pale spot on
segment II, and with triangular basolateral pale patches present but not
visible from above; sterna variable, dark with broad basal pale bands. The
above description has been prepared from the holotype female and 1 paratype
female, both from Sarawak, Malaya.
MALE. Head. Palpus dark, with narrow pale bands at the middle of
segment III, at the base of IV, and at the base and apex of V. Terminalia.
(Figure 75C). Subapical lobe of the basimere well developed, with a blunt
basal rod and 2 hooked rods followed by 3 accessory setae, a broad, striated
leaf, and a slender, gently curved seta; distimere somewhat swollen, with
minute apical annulations on the convex surface; inner division of the phallo-
some with approximately 4 strong teeth, the spiculate portion with a truncate
apex and with a slight extension beyond attachment of the teeth; proctiger (not
illustrated) crowned with a strong tuft of spines, and with 2 or 3 cercal setae
present; basal sternal process strong and well developed.
LARVA. (Figure 75A,B). Head. Antenna with a narrow dark basal
ring and with progressively darker pigmentation beyond insertion of hair 1-A;
head hair 1-C darkly pigmented, bluntly rounded, its length slightly less than
half the distance between the bases of the pair; 4-C single, simple; 5, 6-C
usually bifid, pectinate, subequal in length. Thorax. Integument glabrous;
hairs 1,2,3-P single, pectinate, subequal in length; 4-P with 2 or 3 pectinate
branches; 5,6-P single, pectinate; 7-P with 4 branches, pectinate; 8-P bifid,
pectinate; 14-P single, simple. Abdomen. Integument glabrous; comb
consisting of from 16 to 23 scales arranged in a broad patch (10 to 23 scales
in specimens from Sarawak); individual comb scale elongated, with a promi-
nent apical spine and a Series of fine basolateral spines; siphon broad basally,
tapering characteristically to a narrow apex; siphon index variable, ranging
from 4. 3:1 to 6:1 (5. 9:1 to 7. 9:1 in specimens from Sarawak); 6 pairs of sub-
ventral tufts inserted on the siphon, the subapical pair laterally out of line;
individual tufts 2 to 5 branched, the length of the basal tuft 3 or more times
the width of the siphon at the point of insertion; pecten restricted to approxi-
mately the basal third of the siphon, consisting of from 11 to 14 teeth (12 to
16 teeth in specimens from Sarawak); individual pecten tooth with a promi-
nent apical spine and approximately 10 fine lateral barbs. :
TYPE DATA. Holotype female from Long Tebangan, Fourth Division,
Sarawak, Malaya in the British Museum.
DISTRIBUTION. In THAILAND**, alienus is known from a single col-
lection of fourth stage larvae from Leamngob, Korchang, Tvat. Colless
(1957a) reported the species from SARAWAK, MALAYA, and SINGAPORE.
Fourth stage larvae have been examined in the U. S. National Museum col-
lection from CuChi, SOUTH VIETNAM**, and Tarakan, BORNEO**,
Bram: Genus Culex in Thailand 205
During this study 11 fourth stage larvae have been examined as well
as the holotype female and 2 'morphotypes" with their associated larval and
pupal skins.
TAXONOMIC DISCUSSION. The adult female and the male terminalia
are virtually indistinguishable from pseudovishnui; however, the fourth stage
larva is distinctive and clearly supports Colless' conclusion that this is a
valid species despite its obvious affinity to pseudovishnui and perplexus.
BIOLOGY. Colless (1957a) suggested that the larval habitat is similar
to that of pseudovishnui. The Thailand collection was made from a well, the
South Vietnam collection from a ground pool, and the Borneo collection from
a puddle. Habits of the adults are unknown.
CULEX (CULEX) ANNULUS THEOBALD 1901
(Figures 76, 77, and 78)
Culex annulus Theobald 1901, Mon. Cul. 1: 358 (?); Giles 1902, Handb. 2nd
ed.: 405 (?); Blanchard 1905, Les Moustiques: 293 (?); Leicester
1908, Cul. Malaya: 144 (¢); Theobald 1910, Mon. Cul. 5: 323 (key);
Edwards 1913, Bull. ent. Res. 4: 233 (placed in synonymy with
tritaeniorhynchus).
Culex pseudoinfula Theobald 1911, Tijdschr. Ent. 54: 237 (¢); Colless 1957,
Ann. trop. Med. Parasit. 51: 93 (synonymy).
Culex adelae Baisas 1938, Mon. Bull. Philipp. Bur. Hlth. Serv. 18: 200
(o**, 9, L*, P*); Colless 1957, Ann. trop. Med. Parasit. 51: 93
(synonymy).
Culex (Culex) annulus Theobald: Colless 1957, Ann. trop. Med. Parasit.
51: 93 (o°, 2, L*, revalidated); Delfinado 1966, Mem. Amer. ent. Inst.
1: 130: ({0*, 9, L*),
The adults may not always be conclusively distinguished from several
closely related species of the vishnui subgroup. The fourth stage larva may be
identified by the thoracic and abdominal chaetotaxy, the distinctive comb
scales, and the spiculose thoracic and abdominal integument.
FEMALE. Head. (Figure 76B). Proboscis dark brown with a mod-
erately broad median pale band; palpus similar in color to the proboscis, with
several randomly scattered pale scales at the apex; decumbent scales of the |
vertex pale golden brown at the occiput and becoming white at the orbital line;
erect scales uniformly dull brown, or pale golden at the occiput and dark
brown posterolaterally. Thorax. (Figure 76B,E). Scutum and scutellum
covered with dense, variable, dark brown and golden scales, those of the
fossa usually darker, those of the prescutellar space and scutellum usually
lighter; integument of the pleuron uniformly pale, occasionally with indistinct,
irregular darker areas; patches of dull, white scales on the upper mesepimeron
and the upper and posterior sternopleuron. Wing. (Figure 76C). Dorsal
scales dark brown, but usually with a small patch of pale scales on the
posterior margin of the costa in the area of the humeral cross vein, and oc-
casionally with several other randomly scattered pale scales. Legs. (Figure
76D). Anterior surface of hind femur predominantly pale with a rather nar-
row subapical dark band which extends proximally along the dorsal margin;
hind tibia dark, occasionally with several basal pale scales and a narrow pale
206 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C. (Culex) annulus
Bram: Genus Culex in Thailand 207
apical band; hind tarsus dark, with narrow pale basal bands on tarsomeres
I-IV; fore and mid femora dark, with an apical tuft of pale scales and with
occasional, randomly scattered pale scales; fore and mid tibiae and tarsi
marked, in general, as the hind legs, but with occasional pale or dusky
scales randomly scattered among the dark. Abdomen. (Figure 76F). Terga
dark with variable but usually moderately broad basal white bands on seg-
ments I-VUI; sterna variable, predominantly pale with dark apical bands.
MALE. Head. (Figure 76A). Proboscis with a ventrolateral tuft of
strong setae at approximately the base of the median pale band; palpus dark,
with a broad apical pale band on segment II, a broad median pale band on
segment III, a pale basal band on segment IV, and a basal and apical pale
band on segment V. Terminalia. (Figure T7C). Subapical lobe of the basi-
mere with a blunt basal rod and 2 hooked rods followed by 2 or 3 accessory
setae, a broad striated leaf, and a slender, gently curved seta; distimere
normal in shape, with minute apical annulations on the convex surface; inner
division of the phallosome with 4 strong, darkly pigmented teeth, the spicu-
late portion with a somewhat scalloped apex; proctiger crowned with a strong
tuft of spines and with 2 or 3 cercal setae; basal sternal process well develop-
ed and strongly curved. |
PUPA. (Figure 77A,B). 7
LARVA. (Figure 78). Head. Antenna with a narrow dark basal ring
and somewhat darker beyond insertion of hair 1-A; head hair 1-C darkly
pigmented, robust, tapering to a blunt point, its length about half the distance
between the bases of the pair; 4-C short, simple, single or bifid; 5-C usually
trifid, pectinate; 6-C with 2 or 3 pectinate branches. Thorax. Integument
covered with very minute spicules, particularly on the anterolateral margin;
hairs 1,2,3-P single, pectinate, subequal in length; 4-P bifid, pectinate;
5,6-P single, pectinate; 7-P trifid, pectinate; 8-P bifid, pectinate; 14-P
single, simple. Abdomen. Integument with spicules similar to those of the
thorax; comb consisting of from 25 to 35 scales arranged in a broad, tri-
angular patch; individual comb scale with a prominent apical spine and
fringed laterally with subequal spinules; siphon index variable, ranging from
4:1 to 6:1; 5 or 6 pairs of subventral tufts inserted on the siphon, the ante-
penultimate tuft laterally out of line, and the other tufts inserted at the extreme
margin; individual tufts with 6 or more branches, their length greater than
the width of the siphon at the point of insertion; pecten restricted to the basal
third of the siphon, consisting of approximately 12 rather long teeth; individual
pecten tooth with a strong apical spine and from 4 to 6 strong, lateral barbs,
the apical pecten tooth may have fewer lateral barbs and be somewhat curved.
TYPE DATA. Holotype female of annulus from Hong Kong, China in
the British Museum. Holotype female of pseudoinfula from Pasuruan and
Samarang, Java in the University of Amsterdam Museum, Amsterdam,
Netherlands. The type locality of adelae is Tungkong Manga, San Jose,
Bulacan, Philippines, but the type specimen has been lost.
Figure 76. C. (Culex) annulus. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax;
C, dorsal aspect of the female wing; D, anterior surface of
the female legs; E, dorsal aspect of the female scutum and
scutellum; F, dorsal aspect of the female abdomen.
om
208 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
aT 1), 4s ’
1 Loy me ph lay,
NM ata iy, i
x es 1 Pein Hiates
pupk Wy /
ere Mi) Hh 2 \ Whey?
C.(Culex) annulus
Bram: Genus Culex in Thailand 209
DISTRIBUTION. The only previous THAILAND record for annulus
has been that of Scanlon and Esah (1965) from Doi Pui, Chiang Mai, but the
Species was undoubtedly reported from Thailand prior to 1965 under a dif-
ferent name (see taxonomic discussion below). The species is probably
distributed throughout the country and has been examined from: Chiang Mai,
Chiang Rai, Krung Thep, Lampang, Nakhon Ratchasima, Nan, Pathum Thani,
Phet Buri, Sakon Nakhon, Sara Buri, and Udon Thant.
In addition to Thailand, annulus has been reported from CHINA, JAVA,
SINGAPORE, MALAYA, TAIWAN, NORTH VIETNAM, and the PHILIPPINES.
Specimens have been studied in the U. S. National Museum collection from
Assam, INDIA**, Rangoon, BURMA**, and Phu-tai, Di-An, Soctrang, An-
Khe and Qui-nhon, SOUTH VIETNAM**,
During this study the following specimens have been examined from
Thailand: 78 females and 35 males, 17 with their associated larval and
pupal skins, and 143 larvae.
TAXONOMIC DISCUSSION. From 1913 until revalidated by Colless in
1957, annulus was recognized only as a synonym of tritaeniorhynchus.
During this period it was probably reported from Thailand as the latter
species and, as pointed out by Colless (1957a), also as vishnui.
Although the adult female is very similar to tritaeniorhynchus,
pseudovishnui, and several other members of the vishnui subgroup, the
fourth stage larva is easily distinguished on the basis of the comb scales, the
spiculate thorax, and the multiply branched siphon tufts.
BIOLOGY. Larvae of annulus have been reported from rice fields,
ponds, and ground pools on Taiwan (Lien 1962) and from farm reservoirs and
salt water reservoirs in North Vietnam (Safyanova ef al. 1964). In Thailand,
larvae have also been collected from puddles in dry stream beds, ditches, pot
holes, hoof prints, and on one occasion from a bamboo stump.
Scanlon and Esah (1965) reported abundant night-time human biting
collections of this species in a forest at an elevation of up to 4,500 feet.
Although Lien (1962) reported that this species was especially active at dusk,
Colless (1957b) suggested that the species remains active throughout the
night, but progressively reduced catches reflect a reduction in the immediately
available unfed population, rather than a crepuscular tendency in the species
itself. Lien (1962) has tentatively suggested that annulus may have been
found to harbor first stage filaria larvae of Wuchereria sp., but the collection
probably represents an incidental infection.
CULEX (CULEX) BARRAUDI EDWARDS 1922
(Figure 79)
Culex barraudi Edwards 1922, Indian J. med. Res. 10: 284 (o*); Barraud
1923, Indian J. med. Res. 10: 939 (L*); Barraud 1924, Indian J.
med. Res. 11: 997 (o'*); Senior-White 1927, Spolia. zeylan. 14: 71 (L*).
Figure 77. C. (Culex) annulus. A,B, dorsoventral aspects of the pupa;
C, dorsal aspect of the male terminalia with enlarged
<— insert of the lateral aspect of the inner division of the phallo-
some.
210 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig./8
C. (Culex) annulus
Bram: Genus Culex in Thailand | 211
Culex edwardsi Barraud 1923, Indian J. med. Res. 11: 507 (2); Barraud
1934, Fauna Brit. India, Diptera 5: 397 (o*, 2); Carter and
Wijesundara 1948, Ceylon J. Sci. (B) 23: 148 (L*). NEW
SYNONYMY. :
Culex (Culex) barraudi Edwards: Edwards 1932, in Wytsman, Genera
Insect., fasc. 194: 204 (taxonomy); Barraud 1934, Fauna Brit.
India, Diptera 5: 403 (o‘*, L*); Peters and Dewar 1956, Indian J.
Malar. 10: 46 (o*, L*).
The adults may be identified by the presence of a distinct white line of
scales on the mid and hind femora. The fourth stage larva can be separated
from other members of the vishnui subgroup on the basis of the proximal
prolongation of the stirrup-shaped piece of the siphon.
FEMALE. Head. Proboscis dark brown with a median band of
white scales; palpus similar in color to proboscis, with a dense tuft of pale
scales at the apex of the last segment; decumbent scales of the vertex uni-
formly creamy white; erect scales light brown. Thorax. Scutum covered
with golden brown scales, with some silver scales forming indistinct
patches on the humeral, anterior dorsocentral, antealar areas, and in and
around the prescutellar space; scutellum with scattered white scales; in-
tegument of the pleuron uniformly dark brown; patches of white scales present
on the upper sternopleuron, posterior sternopleuron, anterior mesepimeron,
upper meSepimeron, and the propleuron. Wing. All dorsal wing scales dark
brown. Legs. Anterior surface of the hind femur basically dark, but with a
broad median stripe of white scales which terminates somewhat before the
apex; hind tibia dark, with a narrow median stripe of white scales and with
white scales at the apex; hind tarsomeres I and II with very narrow basal
and apical pale bands; hind tarsomere III with a narrow basal white band, and
IV and V completely dark; mid femur dark, but with a narrow median pale
stripe; mid tibia and tarsus similar to that of the hind leg; fore femur
completely dark; fore tibia with a very narrow basal white band; fore tarsus
Similar to the hind tarsus. Abdomen. Terga dark brown, with moderately
broad basal white bands which extend posteriorly along the lateral margins on
each segment; sterna with the scaling similar to that of the terga.
MALE. Terminalia. (Figure 79C). Subapical lobe of the basimere |
well developed, with 3 hooked rods, 3 rather broad accessory setae, a broad,
striated leaf, anda slender, gently curved seta; distimere normal; inner
division of the phallosome with 4 strong, broad teeth, the spiculate portion
smoothly curved and rather narrow; proctiger crowned with a strong tuft of
Spines and with 3 cercal setae; basal sternal process well developed and
strongly curved.
LARVA. (Figure 79A,B). Head. Antenna with a narrow, dark basal
ring and more darkly pigmented beyond insertion of hair 1-A; head hair 1-C
robust, darkly pigmented, its length approximately half the distance between
the bases of the pair; 4-C single, simple; 5-C with 4 branches, pectinate;
6-C with 3 pectinate branches. Thorax. Integument glabrous; hairs 1,2,3-P
Figure 78. C. (Culex) annulus. Fourth stage larva: dorsoventral view
of the head, thorax and abdomen, and lateral aspect of the
= terminal abdominal segments.
212 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
|
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——— ———
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C. (Culex) barraudi
Bram: Genus Culex in Thailand 213
single, pectinate; 4-P bifid, pectinate; 5,6-P single, pectinate; 7-P trifid,
pectinate; 8-P bifid, pectinate; 14-P single, simple. Abdomen. Integument
glabrous; comb consisting of approximately 45 fan-shaped scales arranged in
a broadly triangular patch; siphon index ranging from 4:1 to 4.5:1, the
siphon tapering gradually to a truncate apex; stirrup-shaped piece at the apex
of the siphon with a prolongation of the ventral cornu directed proximally; 4
pairs of subventral tufts inserted on the siphon, the subapical pair inserted
laterally out of line; individual tufts with from 1 to 4 branches, their length
equal to or less than the width of the siphon at the point of insertion; pecten
consisting of approximately 12 teeth restricted to the basal third of the
siphon; individual pecten tooth with a strong apical spine and from 5 to 9
strong lateral barbs.
TYPE DATA. Holotype male (terminalia slide-mounted) of barvaudi
from Mahdopur, India in the British Museum. Holotype female of edwardsi
from Shillong, Assam, India in the British Museum.
DISTRIBUTION. In THAILAND, barvaudi in known only from Chiang
Mai and Prachuap Khiri Khan. The species has also been recorded from
INDIA, CEYLON, and NEPAL. Specimens have been examined in the U. S.
National Museum from Lahore, EAST PAKISTAN**, and in the British Mu-
seum collection from S. Highlands, Minj., 5,400 ft., T.P.N.G., NEW
GUINEA**,
During this study 2 fourth stage larvae have been examined from Thai-
land. In addition 5 males, 6 females, and 10 larvae have been studied from
areas outside of Thailand.
TAXONOMIC DISCUSSION. The original description of this species by
Edwards and subsequent discussions by Barraud (1923, 1924, 1934) did not
consider ornamentation of the legs, particularly the longitudinal stripes on
the hind and mid femora and tibiae. This feature is also characteristic of
edwardsi, and considering that the male terminalia of both species are iden-
tical, it is concluded that edwardsi and barraudi are conspecific.
BIOLOGY. In Thailand, larvae of barraudi have been collected from
a stream and from a roadside pond, and it is probable that this species is
restricted to rather high elevations. Scanlon and Esah (1965) reported col-
lecting females of this species biting man at between 2, 500 and 4, 500 feet
in a forest environment in Chiang Mai.
CULEX (CULEX) MIMULUS EDWARDS 1915
(Figures 80, 81, and 82)
Culex mimulus Edwards 1915, Bull. ent. Res. 5: 284 (¢,?); Barraud 1923,
Ind. J. med. Res. 10: 942 (L*); Barraud 1924, Ind. J. med. Res.
11: 993 («*); Borel 1926, Arch. Insts. Pasteur Indo-Chine 3-4: 12
(o* 0 Ee),
Figure 79. C. (Culex) barraudi. A, dorsoventral aspect of the head and
prothorax of the fourth stage larva; B, lateral aspect of the
<< terminal abdominal segments of the fourth stage larva; C,
dorsal aspect of the male terminalia with enlarged insert of
the lateral aspect of the inner division of the phallosome.
214 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C. (Culex ) mimulus
Bram: Genus Culex in Thailand 215
Culex mossmani Taylor 1915, Proc. Linn. Soc. N. S. W. 40: 181 (c'*, 2);
Edwards 1924, Bull. ent. Res. 14: 394 (synonymy).
Culex (Culex) mimulus Edwards: Edwards 1922, Ind. J. med. Res. 10: 471
(taxonomy); Barraud 1934, Fauna Brit. India, Diptera 5: 412 (o*, 9,
L*); Edwards 1934, in Barraud, ibid: 450 (key); Bonne-Wepster and
Brug 1937, Geneesk. Tijdschr. Ned.-Ind. 77: 66 (¢, °*); Baisas
1938, Mon. Bull. Philipp. Hlth. Serv. 18: 214 (“*); Bonne-Wepster
and Brug 1939, Geneesk. Tijdschr. Ned.-Ind. 79: 1275 (L*); Lee
1944, Atlas Mosq. Larvae Aust. Reg. : 104 (L*); Bonne-Wepster
1954, Roy. trop. Inst. Amst. Spec. Publ. 111: 127 (o, 9*, L*);
Bonne-Wepster 1954, Doc. Med. Geog. et Trop. 6: 377 (o, 2*, L*);
Safyanova ef al. 1964, Zool. Zhur. 43: 1178 (distribution); Delfinado
1966, Mem. Amer. ent. Inst. 7: 150 (c*, 9).
Culex (Culex) confusus Baisas 1938, Mon. Bull. Philipp. Hlth. Serv. 18:
216 (**, 2); Delfinado 1966, Mem. Amer. ent. Inst. 7: 150 (synonymy).
The adults are readily identified by the presence of distinct patches of
white scales on the wings. The fourth stage larva may be distinguished by
chaetotaxy of the head and thorax, and by features of the siphon.
FEMALE. Head. (Figure 80B). Proboscis dark brown with a median
band of white scales; palpus similar in color to the proboscis with several
randomly placed scales present, particularly on the apical segment; decum-
bent scales of the vertex pale, lighter at the orbital line; erect scales light
golden brown on the occiput, dark brown posteriolaterally. Thorax. (Figure
80B, E). Scutum covered with light bronze-brown scales and having in-
distinct, variable dark and light areas; darker in the area of the fossa, lighter
on the anterior promentory, posterior fossal area, supraalar area, and
around the prescutellar space; integument of the pleuron light brown with
variable and indistinct darker areas present, particularly on the sterno-
pleuron and mesepimeron; dense patches of light scales on the upper sterno-
pleuron, posterior sternopleuron, and anterior mesepimeron. Wing. (Figure
80C). Dorsal wing scales mostly dark, but with variable white patches
usually as illustrated; Rg may or may not be dark basally; Rj usually white
apically, but may occasionally be dark; fork of R2, R3 frequently white, but
may be dark; R2 with or without white scales apically; r-m may or may not
be dark; other minor variations in intensity of pale scaling have also been
noted. Legs. (Figure 80D). Anterior surface of the hind femur mainly
dark but with white scales on the ventral half proximally and with a few white
scales at the apex; hind tibia dark, with a narrow basal white band, a vari-
able stripe of white scales medially, and a narrow apical white band; hind
tarsomeres I-III dark, with narrow basal and apical white bands, tarso-
meres IV and V with basal white bands only, or V without white scales; fore
and mid legs with markings similar to those of the hind legs. Abdomen.
(Figure 80F). Terga dark, with moderately broad convex white basal bands;
Figure 80. C. (Culex) mimulus. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax; C,
pons dorsal aspect of the female wing; D, anterior surface of the
female legs; E, dorsal aspect of the female scutum and
scutellum; F, dorsal aspect of the female abdomen.
216 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C. (Culex) mimulus
Fig. 81
Wy
. “ |
ZZ
Bram: Genus Culex in Thailand 217
sterna primarily pale, but with narrow, dark apical bands.
MALE. Head. (Figure 80A). Palpus dark, with a broad, median
pale band on segment II, a basal white band on segment IV, and basal and
apical pale bands on segment V. Terminalia. (Figure 81C). Subapical lobe
of the basimere well developed, with a blunt basal rod followed by 2 hooked
rods, 3 rather broad accessory setae, a broad leaf, and a slender, gently
curved seta; distimere normal in shape, with or without minute annulations
on the convex surface of the apex; inner division of the phallosome with 4 or
5 very strong, sharply pointed teeth, the spiculate portion also toothed
apically; proctiger crowned with a strong tuft of spines, and with 2 or 3
cercal setae present; basal sternal process well developed and strongly
curved.
PUPA. (Figure 81A, B).
LARVA. (Figure 82). Head. Antenna with a narrow dark basal
band and progressively darker beyond insertion of hair 1-A; head hair 1-C
robust, strongly pigmented, gradually tapering to a sharp point, its length
1/3 to 1/2 the distance between the bases of the pair; 4-C usually single, oc-
casionally double, simple; 5-C with from 3 to 6 branches, pectinate; 6-C
usually bifid (occasionally trifid), pectinate. Thorax. Integument glabrous;
hairs 1,2,3-P single, pectinate, subequal in length; 4-P usually single
(rarely bifid), pectinate; 5,6-P single, pectinate; 7-P trifid, pectinate; 8-P
bifid, pectinate; 14-P single, simple. Abdomen. Integument glabrous;
comb variable, consisting of from 20 to 45 scales arranged in a triangular
patch; individual comb scale fan-shaped, but with the median apical spine
frequently slightly longer than the lateral ones; siphon index extremely
variable, ranging from 5:1 to 12:1, if the siphon is short it is usually
truncate apically, if long it is gradually tapering; 5 to 7 pairs of subventral
tufts inserted on the siphon, the subapical tuft laterally out of line; individual
tufts with from 1 to 4 branches, their length usually less than the width of the
siphon at the point of insertion; pecten consisting of approximately 12 to 15
teeth restricted to the basal 1/5 to 1/3 of the siphon depending on the siphon
length; individual pecten tooth with a prominent apical spine and from 4 to 6
lateral barbs, the apical pecten tooth may be somewhat Separated from the
rest of the pecten and gently curved.
TYPE DATA. Holotype male (terminalia slide-mounted) of mimulus
from ''Kuching?, Sarawak, Borneo" in the British Museum. Type of mossmani
from Mossman, Queensland, Australia in the University of Sydney, New
South Wales. The type locality of confusus is Balabac, Balabac Island,
Palawan, Philippines, but the type specimen has been lost.
DISTRIBUTION. In THAILAND, specimens of mimulus have been col-
lected from: Chiang Mai, Chiang Rai, Chon Buri, Kampong Pate, Kanchana-
buri, Lampang, Lamphun, Mae Hong Son, Nakhon Nayok, Nakhon Ratchasima,
Narathiwat, Ranong, Sara Buri, Satun, Songkhla, Surat Thani, Tak, Trang,
and T'vat. This species has also been reported from MALAYA, INDONESIA,
INDOCHINA, CHINA, BURMA, NEPAL, INDIA, CEYLON, NEW GUINEA,
AUSTRALIA, and the PHILIPPINES.
Figure 81. C. (Culex) mimulus. A,B, dorsoventral aspects of the pupa;
“ C, dorsal aspect of the male terminalia with enlarged insert
of the lateral aspect of the inner division of the phallosome.
218 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.82
ENG
eel 1 3 ‘
ees
|
C. (Culex) mimulus
Bram: Genus Culex in Thailand 219
During this study the following specimens from Thailand have been
examined: 172 females and 143 males, 60 with their associated larval and
pupal skins, and 296 larvae.
TAXONOMIC DISCUSSION. When extensive series of this species
were originally examined, it was felt that several anatomical and colorational
characters were indicative of species or subspecies recognition. In the male
terminalia, the distimere may be minutely annulate on the convex surface
apically or smooth; in the larva, the siphon may be relatively long and nar-
row, or short and broad; however, no correlation could be established between
these 2 characters. Other characters, seemingly associated with the long
siphon condition, include extensive white scaling on the wing veins of the
adult female (involving vein Rs and the fork of R2 and Rg) and the degree of
basal white banding on the abdominal terga. Again, correlation could not be
established between larval anatomy and adult ornamentation and, in fact, so
much variation and integradation occurred that even infraspecific categories
could not be established. In general, populations from northern Thailand
demonstrate a short, stout siphon, no white scales on wing vein Rg or the
fork of R2 and R3, extensive abdominal banding, and a smooth distimere;
contrasting this are specimens from southern Thailand, Indonesia, and the
Philippines which exhibit a long, narrow siphon, extensive white scale pat-
terns on the wing, sparse banding on abdominal terga VI to VIII, and an an-
nulate distimere (the type male being included among the latter group). How-
ever, integradation and mixing of characters is common and definite zoo-
geographical forms cannot be established. It is, therefore, concluded that
mimulus is a plastic species which, although apparently undergoing specia-
tion, cannot be validly divided into specific or infraspecific taxa at the present
time. Future collections may supply sufficient biological data so that mean-
ingful correlations between anatomy and ecology can be established.
The adult of mimulus may be separated from its close relative,
mimeticus, by having the first pale costal spot (at middle of wing) extending
over vein R, whereas in mimeticus this spot extends only to the subcosta; in
the fourth stage larva the comb scales of mimeticus exhibit a strongly
developed, long and broad median distal spine, whereas this spine in
mimulus is, at most, only slightly longer than the lateral ones.
BIOLOGY. In Thailand, larvae have been collected from a wide
variety of ground water sources as well as from sumps, hoofprints, stream
pools, and rock holes. Larvae have additionally been reported from sphagnum
pools, swamps covered with vegetation, tree holes, and broken bamboo
Stumps. Although adults have been collected in houses, nothing is known of
the feeding habits; however, man is certainly not a primary, or even
secondary host.
Figure 82. C. (Culex) mimulus. Fourth stage larva: dorsoventral view
of the head, thorax and abdomen, and lateral aspect of the
— terminal abdominal segments.
vol. 2, no. 1, 1967
220
—>
Contrib. Amer. Ent. Inst.,
iy
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|
JTL. HWANG
C. (Culex) perplexus
Bram: Genus Culex in Thailand 221
CULEX (CULEX) PERPLEXUS LEICESTER 1908
(Figure 83)
Culex perplexus Leicester 1908, Cul. Malaya: 150 (, 2); Edwards 1917,
Bull. ent. Res. 7:-225 (placed in synonymy with vishnuz); Colless
1957, Ann. trop. Med. Parasit. 51: 95 (“,?, L*, revalidated).
The adults may be recognized by the unbanded abdominal terga in
some specimens; other specimens with banded abdominal terga cannot be
conclusively separated from Several closely related members of the vishnui
subgroup. The fourth stage larva is characterized principally by the
distinctive comb, pecten, and siphon shape.
FEMALE. Head. Proboscis dark brown with a rather narrow median
band of light scales; palpus similar in color to the proboscis, the apical seg-
ment tipped with a few white scales; decumbent scales of the vertex light
golden brown, becoming white at the orbital line; erect scales pale golden
brown medially and dark brown posteriolaterally. Thovax. Scutum covered
with dense, narrow pale golden scales anteriorly and dark brown scales from
the supraalar area posteriorly, except for a few light scales surrounding the
prescutellar space and on the scutellum; integument of the pleuron uniformly
dark brown, with patches of dull white scales on the upper sternopleuron,
posterior sternopleuron, and anterior mesepimeron. Wing. All dorsal wing
scales uniformly bronze-brown. Legs. Anterior surface of the hind femur
with rather pale brown scales basally and toward the apical fourth, with dark
scales at the apex and along the dorsal margin, and with a tuft of a few pale
scales at the extreme apex; hind tibia dark but with a few pale scales at the
apex; hind tarsus dark, with very narrow pale basal bands on tarsomeres I-
IiI; fore and mid femora without pale scales or with only a narrow basal band
of pale scales; fore and mid tibiae and tarsi marked as the hind legs, but
tarsal markings reduced. Abdomen. Terga dark brown, in some specimens
without pale basal bands, in others with narrow convex basal bands; sterna
dark brown, with rather broad pale basal bands.
MALE. Head. Palpus with a broad median pale band on segment III,
a basal white band on segment IV, and basal and apical bands on segment V.
Terminalia. (Figure 83B). Subapical lobe of the basimere with a blunt basal
rod followed by 2 hooked rods, 4 accessory setae, a broad leaf, and a slender,
gently curved seta; inner division of the phallosome with 4 strong teeth, the
Sspiculate portion somewhat scalloped apically; proctiger crowned with a
strong tuft of spines and with 2 or 3 cercal setae; basal sternal process well
developed and strongly curved.
LARVA. (Figure 83A,C). Head. Antenna with a narrow dark basal
ring and frequently darker beyond insertion of hair 1-A; head hair 1-C robust,
darkly pigmented, gradually tapering to a sharp point and frequently with
from 1 to 3 accessory lateral spicules; 4-C. single, simple; 5-C usually with
eee emer emcee mentinenmmmememameare meee ee
Figure 83. C. (Culex) perplexus. <A, dorsoventral aspect of the head and
prothorax of the fourth stage larva; B, dorsal aspect of the
<iGiene male terminalia with enlarged insert of the lateral aspect of the
inner division of the phallosome; C, lateral aspect of the
terminal abdominal segments of the fourth stage larva.
222 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
4 or 5 branches, pectinate; 6-C usually bifid, pectinate. Thorax. Integu-
ment glabrous; hairs 1,2,3-P single, pectinate, subequal in length; 4-P
usually bifid, pectinate; 5,6-P single, pectinate; 7-P trifid, pectinate; 8-P
bifid, pectinate; 14-P single, simple. Abdomen. Integument glabrous; comb
consisting of from 12 to 20 scales arranged in 2 or 3 irregular rows; indivi-
dual comb scale elongate, strong, with a prominent distal spine fringed with
fine lateral spicules; siphon index variable, ranging from 5.5:1 to 8.0:1, the
siphon rather narrow and usually with a gentle distal curvature; 5 to 7 ir-
regular pairs of subventral tufts inserted on the siphon, the subapical tuft
inserted laterally out of line; individual tufts with from 3 to 5 simple branches,
their length greater than the width of the siphon at the point of insertion;
pecten variable, consisting of from 8 to 12 teeth restricted to the basal fourth
of the siphon; individual pecten tooth robust, with a prominent apical spine
and a fringe of from 10 to 15 lateral barbs; the distal 2 or 3 pecten teeth more
widely separated from the rest of the pecten, these teeth usually without
lateral barbs or, at least, with a reduced number.
TYPE DATA. Lectotype hereby designated: syntype female, ''Kuala
Lumpur, F. M. S., G. F. Leicester, 1914-274, pupae from large stagnant
pool filled with floating water weed near houses, 12/3/03", in the British
Museum.
DISTRIBUTION. In THAILAND**, perplexus has been collected from:
Phet Buri, Prachin Buri, Nakhon Ratchasima, and Surat Thani (?). Other
known localities include the type locality of Kuala Lumpur, MALAYA and
SINGAPORE.
The following specimens from Thailand have been examined during
this study: 15 females and 10 males, 6 with their associated larval and
pupal skins, and 45 larvae.
TAXONOMIC DISCUSSION. The concept of perplexus as here presented
conforms to the description by Colless (1957a). The adult female of perplexus
demonstrates extremely close affinity to pseudovishnui, and in specimens
which exhibit convex basal abdominal bands, the 2 species cannot be distin-
guished with any degree of certainty. The male terminalia of the 2 species
are also extremely similar. In the fourth stage larva the comb, pecten, and
shape of the siphon of perplexus is distinctive.
BIOLOGY. Larvae of perplexus have been collected in Thailand from
a sump with heavy pollution and light vegetation, from a Swamp, and from
stream pools and stream margins during the months of January and February.
Colless (1957c) reported perplexus as one of the principal culicine species
found in hyacinth ponds at Singapore. Biology of the adults is unknown.
CULEX (CULEX) PSEUDOVISHNUI COLLESS 1957
(Figure 84)
Culex pseudovishnui Colless 1957, Ann. trop. Med. Parasit. 51: 88 (o*, 9, L*).
Figure 84. C. (Culex) pseudovishnui. A, dorsoventral aspect of the head
and prothorax of the fourth stage larva; B, lateral aspect of
the terminal abdominal segments of the fourth stage larva; C,
dorsal aspect of the male terminalia with enlarged insert of
the lateral aspect of the inner division of the phallosome.
Bram: Genus Culex in Thailand 223
Fig.84
K ie
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XN SX
C. (Culex ) pseudovishnui
224 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
The adults are most difficult to separate from Several other members
of the vishnui subgroup but, in general, may be recognized by the banding of
the hind femur and the color patterns of the erect scales of the vertex. The
fourth stage larva is characterized by the distinctive comb, pecten, and
siphon. :
FEMALE. Head. Proboscis dark brown with a moderately broad
median band of pale scales; palpus similar in color to the proboscis, oc-
casionally tipped with several randomly placed pale scales; decumbent scales
of the vertex pale golden brown, becoming white at the orbital line; erect
scales pale golden brown at the occiput and dark brown posteriolaterally.
Thorax. Scutum with a variable grouping of brown, pale golden, and white
scales, darker at the fossa and along the acrostichal and posterior dorso-
central lines, lighter on the lateral prescutal and supraalar areas; integu-
ment of the pleuron uniformly light brown, but occasionally with some ir-
regular darker areas; patches of white scales present on the upper and
posterior sternopleuron and the upper mesepimeron. Wing. All dorsal wing
scales dark brown. Legs. Anterior surface of the hind femur mainly pale,
with a broad dark subapical band which extends proximally to form a dark
stripe along the dorsal margin; hind tibia dark with a narrow apical band of
pale scales and a poorly defined stripe posteriorly; hind tarsus dark, with
narrow pale basal bands on tarsomeres I-III, occasionally on tarsomeres IV
and V, and a narrow apical pale band on tarsomere J and often on II; anterior
surface of the fore and mid femora usually dark with narrow basal and apical
pale bands; fore and mid tibiae and tarsi usually marked as the hind legs.
Abdomen. Terga dark with narrow pale basal bands on segments III-VII,
terga II usually with a central basal pale patch; terga II-VII with large baso-
lateral pale spots not visible from above; sterna predominantly pale with
variable dark apical bands.
MALE. Head. Palpus dark, segment III with a narrow pale basal
band and a broad median pale band, IV with a basal white band, and V with
basal and usually apical white bands. Terminalia. (Figure 84C). Subapical
lobe of the basimere well developed, with a blunt basal rod and 2 hooked rods
followed by 3 or 4 accessory setae (one of which may be very broad), a broad,
striated leaf, and a seta; distimere normal in shape, with fine annulations
on the convex surface at the apex; inner division of the phallosome with 3 or 4
strong teeth, the spiculate portion very finely toothed or scalloped apically;
proctiger crowned with a strong tuft of spines, and with 2 or 3 cercal setae
present; basal sternal process well developed and strongly curved.
LARVA. (Figure 84A,B). Head. Antenna with a narrow, dark basal
ring and progressively darker beyond insertion of hair 1-A; head hair 1-C
strongly pigmented, robust, gradually tapering to a sharp point, its length
about half the distance between the bases of the pair; 4-C single or branched,
simple; 5-C with 3 or 4 branches, pectinate; 6-C with 2 or 3 branches, pecti-
nate. Thorax. Integument usually glabrous, rarely covered with a random
pattern of minute spicules; hairs 1,2,3-P single, pectinate, subequal in
length; 4-P most frequently bifid or occasionally trifid, pectinate, its length
about 3/4 the length of 3-P; in some specimens 4-P is weak and short and
possesses from 5 to 10 distinct branches; 5,6-P single, pectinate; 7-P tri-
fid, pectinate; 8-P bifid, pectinate; 14-P single, simple. Abdomen. Integu-
ment similar to that of the thorax; comb consisting of from 5 to 13 scales ar-
ranged in an irregular row; individual comb scale elongate, with an extremely
Bram: Genus Culex in Thailand 225
prominent apical spine and with inconspicuous lateral spicules; siphon index
variable, ranging from 4. 5:1 to 8.5:1; 5 to 7 irregular pairs of subventral
tufts inserted on the siphon as well as from 1 to 3 irregular pairs of lateral
tufts; individual tufts with.from 2 to 6 simple branches, their length greater
than the width of the siphon at the point of insertion; pecten consisting of from
8 to 11 teeth restricted to the basal third of the siphon; individual pecten
tooth with a prominent apical spine and from 6 to 12 basolateral barbs.
TYPE DATA. Holotype female with associated larval and pupal skins
from Singapore in the British Museum.
DISTRIBUTION. In THAILAND, pseudovishnui is probably widely
distributed throughout the country and has been studied from: Chiang Mai,
Chiang Rai, Chon Buri, Krung Thep, Lampang, Mae Hong Son, Nakhon Nayok,
Nakhon Ratchasima, Nan, Narathiwat, Patum Thani, Phet Buri, Prachuap
khiri Khan, Ranong, Rayong, Surat Thani, Trang, Udon Thani, and Yala, .
The species is also widely distributed throughout the ORIENTAL REGION.
During this study the following specimens have been examined from
Thailand: 104 females and 18 males, 10 with their associated larval and
pupal skins, and 96 larvae.
TAXONOMIC DISCUSSION. This species has been most frequcnily
misidentified as vishnui. During this study an attempt was made to correlate
anatomical features of the head hairs, comb, pecten, and siphon with distri-
butional patterns. In no instance could correlations be established, although
in general it appears that the condition in which thoracic hair 4-P is bifid
and stout is most frequently found in the more northern areas (Thailand,
Taiwan, etc.) and the multiple-branched, fine condition is seen in the more
southern areas (Indonesia, Malaya, Philippines), although both conditions
are probably found throughout the range of the species.
BIOLOGY. In Thailand, larvae of pseudovishnui have been collected
from a wide variety of ground water habitats including ditches, rice fields,
ponds, streams, and sumps. Colless (1959b) reported that the adult females
feed primarily on birds and pig, and to a lesser degree on ox, dog, horse,
and man. Scanlon and Esah (1965) collected females biting man at elevations
ereater than 4,500 feet on Doi Pui, Chiang Mai, Thailand.
CULEX (CULEX) TRITAENIORHYNCHUS GILES 1901
(Figures 85, 86, and 87)
Culex tritaeniorhynchus Giles 1901, J. Bombay nat. Hist. Soc. 13: 606
(adult); Theobald 1901, Mon. Cul. 1: 364 (c'*, ?*); Giles 1902, Handb. ,
and ed.: 401 (<, 9); Edwards 1913, Bull. ent. Res. 4: 233 (taxonomy);
Edwards 1917, Bull. ent. Res. 7: 224 (“, 2); Edwards 1921, Bull.
ent. Res. 12: 339 (L); Macfie and Ingram 1922, Ann. trop. Med.
Parasit. 16: 180 (?*); Barraud 1923, Ind. J. med. Res. 10: 940 (L*);
Barraud 1924, Ind. J. med. Res. 11: 995 (o*, 2); Borel 1930,
Monogr. Coll. Soc. Pat. exot. 3: 336 (o*, 2, L*): Ho 1931, Bull. Fan.
Inst. Biol. , Peking 2: 158 (o*, 9).
Culex birot Theobald 1905, Ann. hist.-nat. Mus. hung. 3: 82 (“*,?); Edwards
1913, Bull. ent. Res. 4: 233 (synonymy).
Culex summorosus Dyar 1920, Insec. Inscit. menst. 8: 180 (<); Baisas 1938,
Mon. Bull. Philipp. Hlth. Serv. 18: 196 (o*, 9*, P*, L*); Colless 1957,
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
226
Bram: Genus Culex in Thailand 227
Ann. trop. Med. Parasit. 51: 98 («*,?, L*, as ssp. of
tritaeniorhynchus) ; Delfinado 1966, Mem. Amer. ent. Inst. 7: 153
(o*,9,P,L*, as ssp. of tritaeniorhynchus). NEW SYNONYMY.
Culex tritaeniorhynchus var. siamensis Barraud and Christophers 1931,
Rec. Malar. Surv. India 2: 283 (o); Brug 1934, Bull. ent. Res. 25:
515 (o); Bonne-Wepster and Brug 1937, Geneesk. Tijdschr. Ned. -
Ind. 77: 72 (o,?*); Bonne-Wepster and Brug 1939, Geneesk.
Tijdschr. Ned.-Ind. 79: 1270 (L*); Lee 1944, Atlas Mosq. Larvae
Aust. Reg.: 105 (L*); Bonne-Wepster 1954, Roy. trop. Inst. Amst.,
spec. publ. 111: 125 (o, 9*, L*); Mattingly 1956, Proc. R. ent. Soc.
Lond. (A) 31: 37 (lectotype designation); Colless 1957, Ann. trop.
Med. Parasit. 51: 98 (as synonym of tvitaeniorhynchus summorosus).
Culex (Culex) tritaeniorhynchus Giles: Edwards 1932, in Wytsman, Gen.
Insect. , fasc. 194: 204 (taxonomy); Barraud 1934, Fauna Brit. India,
Diptera 5: 404 (o*, 9, L*); Li and Wu 1935, Yearb. Bur. Ent. Chekiang
4: 101 (L*); Stackelberg 1937, Faune de 1' URSS, Ins. Dipt. 3: 235
(o*); Edwards 1941, Mosq. Ethiop. Reg. 3: 299 (<, 2); Bohart 1945,
Navmed 580: 80 (*, L); Bohart and Ingram 1946, Navmed 1055: 81
(o*, 2, L*); LaCasse and Yamaguti 1950, Mosq. Fauna Japan
Korea: 230 (o*, 9*, P*, L*); Monchadskii 1951, Moscow zool. Inst.
Acad. Nauk SSSR 37: 277 (L*); Iyengar and Menon 1955, Bull. ent.
Res. 46: 6 (L*); Khattat 1955, Bull. end. Dis. 1: 165 (L*); Bohart
1956, Ins. Micronesia 12: 83 (c'*, 2, L*); Ovazza, Hamon, and Nori
1956, Soc. Path. exot. Bull. 49: 169 (“*); Ma and Feng 1956, Acta
ent. Sinica 6: 172 (L*); Hara 1957, Jap. J. exp. Med. 27: 56 (?*);
Senevet and Andarelli 1959, Encycl. ent. (A) 37: 142 (o,?, L, P);
Foote and Cook 1959, USDA Agric. Handb. 152: 140 (biology).
The adult female is most difficult to distinguish from several other
members of the subgroup, but may usually be recognized by the rather small
size, the brown scaling of the scutum, the accessory pale scaling on the
ventral surface of the proboscis; the long, filamentous teeth of the bucco-
pharyngeal armature are diagnostic. In the adult male, the terminalia is also
Similar to other members of the subgroup, but may be recognized to some
degree by the structure of the inner division of the phallosome. The fourth
stage larva is characterized by the thoracic chaetotaxy, the shape of the comb
scales, and by anatomical features of the siphon.
FEMALE. A rather small species with overall dull appearance. Head.
(Figure 85B). Proboscis dark brown, with a moderate sized median band of
pale scales and usually with accessory patches of pale scales on the ventral
surface proximal to the band (this character is extremely variable and may
range from no pale scales to a condition in which virtually the entire ventral
surface of the proboscis proximal to the band is pale); a few randomly scattered
Figure 85. C. (Culex) tritaeniorhynchus. Adult habitus: A, lateral
aspect of the male head; B, lateral aspect of the female head
— and thorax; C, dorsal aspect of the female buccophar yngeal
armature; D, dorsal aspect of the female wing; E, anterior
surface of the female legs; F, dorsal aspect of the female
scutum and scutellum; G, dorasl aspect of the female abdomen.
228 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
pale scales may occasionally be present on the dorsal surface of the pro-
boscis; palpus similar in color to the proboscis, the apex occasionally ex-
hibiting several randomly placed scales; decumbent scales of the vertex
golden brown, becoming somewhat paler at the orbital line; erect scales.
dark brown; teeth of the buccopharyngeal armature very long and filamentous
(figure g5c)!. Thorax. (Figure 85B, F). Scutum almost uniformly covered
with dense, bronze-brown scales, but with a few paler scales on the supra-
alar and prescutellar areas; integument of the pleuron usually pale brown,
occasionally with variable and irregular darker and paler areas; patches of
dull white scales present on the anterior mesepimeron, and upper and lower
sternopleuron. Wing. (Figure 85D). Legs. (Figure 85E). Anterior surface
of the hind femur predominantly pale with a narrow dark apical band which
frequently extends proximally along the dorsal border; hind tibia dark, with a
narrow basal pale band; hind tarsus dark with narrow, pale basal bands on
tarsomeres I-V; fore and mid femora variable, usually darker than the hind
femur; fore and mid tibiae and tarsi variable, but in general similar to the
hind legs. Abdomen. (Figure 85G). Terga dark with variable, usually nar-
row, convex pale basal bands (these bands are occasionally represented by
only a few pale scales); sterna variable, predominantly pale, usually with
rather broad apical dark bands.
MALE. Head. (Figure 85A). Proboscis with a rather narrow median
pale band, and with a small ventrolateral tuft of setae at approximately the
base of the median band; palpus dark, with a narrow basal and usually a broad
median pale band on segment III and with narrow basal bands on segments IV
and V. Terminalia. (Figure 86C). Subapical lobe of the basimere well
developed, with a blunt basal rod followed by 2 subequal, hooked rods, 4 ac-
cessory setae (one of which is quite broad and long), a broad, striated leaf,
and a slender, gently curved seta; distimere normal in shape, with minute
annulations on the convex surface at the apex; inner division of the phallo-
some usually with 4 strong, recurved teeth, the spiculate portion with a
slightly expanded apex projecting somewhat beyond the attachment of the
teeth; proctiger crowned with a strong tuft of spines and with 2 or 3 cercal
setae; basal sternal process well developed and strongly recurved.
PUPA. (Figure 86A,B).
LARVA. (Figure 87). Head. Antenna with a narrow, dark basal
ring and somewhat darker beyond insertion of hair 1-A; head hair 1-C strongly
pigmented, gradually tapering to a sharp point, its length slightly greater
than half the distance between the bases of the pair; 4-C single, simple; 5-C
with 3 or 4 pectinate branches; 6-C with 2 or 3 pectinate branches. Thorax.
: de Meillon (1967 - personal communication) has examined specimens of
tritaeniorhynchus from: Ramsar, Iran; Chinhae, Korea; Pakistan; Honshu,
Japan; Karwar, India; Port Lewis, Mauiritius; and Al Khabar, Saudi Arabia.
In each instance the anatomy of the buccopharyngeal armature has reamined
consistent.
Figure 86. § C. (Culex) tritaeniorhynchus. A,B, dorsoventral aspects of
the pupa; C, dorsal aspect of the male terminalia with enlarged
insert of the lateral aspect of the inner division of the phallo-
some. a
Bram: Genus Culex in Thailand
\\ as
YESS
SASS \\
C. (Culex) tritaeniorhynchus
ve
230 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Integument glabrous; hairs 1,2,3-P single, pectinate, subequal in length; 4-P
usually bifid, pectinate; 5,6-P single, pectinate; 7-P trifid, pectinate; 8-P
bifid, pectinate; 14-P single, simple. Abdomen. Integument glabrous; 1-X
with 3 or 4 short branches; comb variable, consisting of from 25 to 40 evenly
fringed, fan-shaped scales arranged in a triangular patch of 3 irregular rows;
Siphon index variable, ranging from approximately 5:1 to 9.5:1 (average,
7.5:1), the siphon evenly tapering from base to apex; stirrup-shaped piece
without a prolongation of the ventral cornu; from 5 to 7 (usually 7) pairs of
subventral tufts inserted on the siphon, the subapical tuft inserted laterally
out of line; individual tufts with from 2 to 5 branches, their length slightly
greater than the width of the siphon at the point of insertion; pecten consisting
of from 9 to 15 teeth restricted to the basal third of the siphon or less; indi-
vidual pecten tooth with a prominent apical spine and approximately 6 lateral
barbs.
TYPE DATA. Holotype female of tritaeniorhynchus from Travancore,
India in the British Museum. Syntypes male and female of bivoi from Bombay,
India in the Magyar Nemzeti Muzeum, Budapest, Hungary. Holotype male
(terminalia slide-mounted) of swmmorosus from Los Banos, Luzon, Philip-
pines in the U. S. National Museum. Lectotype male (terminalia slide-
mounted) of tvitaeniorhynchus var. siamensis from Chiang Mai, Thailand in
the British Museum.
DISTRIBUTION. This species is undoubtedly widely distributed through-
out THAILAND and during this study specimens have been examined from:
Ayutthaya, Chanthaburi, Chiang Mai, Chiang Rai, Chon Buvi, Lampang,
Lamphun, Lop Buri, Nakhon Nayok, Nakhon Ratchasima, Nan, Nonthabuyi,
Pathum Thani, Phet Buri, Prachuap Khiri Khan, Ranong, Rayong, Roi Et,
Samut Prakan, Sara Buri, Songkhla, Surat Thani, Thon Buri, Trang, Udon
Thani, and Yala.
C. tritaeniorhynchus has been reported from: INDIA, CEYLON,
MALDIVE ISLANDS, MALAGASY, MOCAMBIQUE, ZANZIBAR, KENYA,
UBANGI-SHARI, NIGERIA, DAHOMEY, TOGO, GOLD COAST, GAMBIA,
SENEGAL, EGYPT, ARABIAN PENINSULA, ISRAEL, LEBANON, SYRIA,
TURKEY, IRAQ, IRAN, TRANSCAUCASIA, and TURKMEN §S. S. R. It has
also been reported as tritaeniorhynchus ssp. summorosus from: PHILIP-
PINES, TAIWAN, OGASAWARA GUNTO, RYUKYU-RETTO, JAPAN, KOREA,
MANCHURIA, CHINA, INDO-CHINA, INDONESIA, MALAYA, INDIA, and
MARITIME PROVINCE U. S. S. R. Specimens have been examined in the
British Museum collection from Rangoon, BURMA**,
During this study, the following specimens were examined from Thai-
land: 135 females and 83 males, 10 with their associated larval and pupal
skins, as well as 400 larvae. In addition, material has been examined from:
Malaya, Okinawa, Philippines, Borneo, South Vietnam, Pakistan, Iran,
Saudi Arabia, Taiwan, Japan, Korea, India, Iraq, and Egypt.
TAXONOMIC DISCUSSION. Colless (1957a) formally recognized
summorosus aS a Subspecies of tritaeniorhynchus, although he did so with
reservations. Two anatomical features were cited as the primary differ entiat-
Figure 87. C. (Culex) tritaeniorhynchus. Fourth stage larva: dorsoventral
view of the head, thorax and abdomen, and lateral aspect of the
terminal abdominal segments.
Bram: Genus Culex in Thailand
C. (Culex) tritaeniorhynchus
231
Fig.87
Pa x
V =
= ls ——_———o
= —
~ CG
——
232 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
ing characteristics for the 2 subspecies (i.e., ¢. tritaeniorhynchus and t.
summorosus). The first of these is the form and structure of the lateral
plate of the phallosome of the male terminalia: in ¢vifaeniorhynchus the
teeth are somewhat weakly developed and only gently curved, where in
summorosus they are strongly developed, considerably longer, and sharply
recurved. The second feature is the general length and shape of the larval
siphon; in tviiaeniorhynchus the siphon is rather short, the sides parallel,
and the apex truncate, and in summorosus the siphon is slenderer (particularly
towards the apex) and considerably longer.
Variation exists in both structures, and material collected from Thai-
land suggests integration of these characters; however, in the material from
Thailand both extremes have been Seen as well as a large number of inter-
mediate forms. Variation and integradation appears to be most pronounced
in the larva, with the greatest majority of specimens from Thailand conform-
ing to the summorosus form sensu Colless. A form with an extremely long
siphon (See Barraud 1934 and Colless 1957a) has been found principally in
southern Thailand (from Bangkok southward). The form with the short siphon,
or tritaeniorhynchus sensu Colless, has been studied from Chiang Mai,
Songkhla, and Roi Et (changwats located in the northern, penninsular, and
eastern parts of the country). A limited amount of material was available for
study from India, Irag, and Egypt, but in each case these specimens exhibited
the short siphon. Consideration of variation within the two primary differ entiat-
ing characteristics strongly suggests, at the present time, that friiaeniorhynchus
represents a plastic species with clinal variation in certain anatomical features,
but is not amenable to a strictly subspecific interpretation. On the basis of
data from specimens collected in Thailand, it is concluded that summorosus
should be recognized as a synonym of tritaeniorhynchus until additional
anatomical, biological, and vector potential data are available for the species
throughout its range.
Due to the difficulty in conclusively recognizing some adults of
tritaeniorhynchus, it has been necessary to resort to the buccopharyngeal arma-
ture of the female as a supplementary character. In tritaeniorhynchus the
teeth of the buccopharyngeal armature are quite long and filamentous, whereas
in the other closely related members of the subgroup studied (annulus,
perplexus, pseudovishnui, and whitei, as well as both species of the sitiens |
subgroup) these teeth are short and acuminate.
BIOLOGY. In Thailand, larvae have been collected from a wide variety
of temporary and semipermanent ground water habitats and have also been
collected from streams and swamps. Colless (1957c) found this to be the most
common species in hyacinth ponds and suggested that almost any sunlit
permanent fresh water pond containing some vegetation would be particularly
favorable. Larvae were also reported from tidal marshes of low salinity,
transient pools with emergent vegetation, obstructed streams, and oc-
casionally from tins or durms sunk into the earth and showing features in
common with ground pools. In Japan, LaCasse and Yamaguti (1950) reported
that the larvae were collected from fresh water habitats early in the season
and as the season advanced, the diversity of situations in which larvae were
collected broadened to include practically all habitats (except container habi-
tats, tree holes, and some unspecified specialized habitats).
The adult females feed primarily on cow and pig, but will occasionally
feed on birds and man (Colless 1959b). Colless (1959c) concluded that in the
Bram: Genus Culex in Thailand 233
absence of cattle, the species is attracted to man but is slow to feed, whereas
in the presence of cattle, man is almost completely ignored. Flemings
(1959) noted that females will bite black crowned night herons (a host of
Japanese encephalitis virus) at all levels of elevation up to 50 ft., but the
biting rate was overwhelmingly higher at ground level and decreased with
altitude. Scanlon and Esah (1965) collected females biting man at elevations
up to 4,500 ft. on a mountain in Chiang Mai. Nakao (1959) has suggested
that tritaeniorhynchus may be an indoor resting species.
This species has been incriminated as a vector of Japanese encephalitis
virus (Tigertt and Hammon 1950, and other authors) and is generally regarded
as the principle vector in the Oriental region. Dengue virus has also been
isolated from tritaeniorhynchus (Rudnick and Hammon 1961), but no epidemio-
logical evidence is available to suggest that it serves any role as a natural
vector. Evaluation of the role tvitaeniorhynchus plays in the transmission of
arthropod-borne viruses is complicated by the afore mentioned difficulties in
conclusively recognizing the females. Caution must, therefore, be exercised
in the collection and interpretation of data relating to this and other species
of the vishnui subgroup.
CULEX (CULEX) WHITEI BARRAUD 1923
(Figure 88)
Culex whitet Barraud 1923, Ind. J. med. Res. 11: 508 (%, 2); Barraud 1924,
Ind. J. med. Res. 11: 998 (*).
Culex (Culex) whitei Barraud: Edwards 1932, in Wytsman, Gen. Insect., fasc.
194: 205 (taxonomy); Barraud 1934, Fauna Brit. Ind., Diptera 5: 402
(o**, L); Peters and Dewar 1956, Ind. J. Malar. 10: 46 (L*).
The adult female cannot be conclusively distinguished from several
other members of the vishnui subgroup. The male terminalia exhibits
distinctive setae on the subapical lobe of the basimere and possesses 3 or
more strong teeth on the outer division of the phallosome. The fourth stage
larva is distinctive in the shape of the comb scale, and the chaetotaxy of the
head and siphon.
FEMALE. Head. Proboscis dark brown ae a moderately sized
median band of pale scales; palpus dark brown, without pale scales; decum-
bent scales of the vertex pale brown at the occiput, becoming almost white
at the orbital line; erect scales pale golden to light orange at the occiput and
dark brown posteriolaterally. Thorax. Scutum covered with a variable, ir-
regular pattern of pale golden and dark brown scales, usually darkest on the
fossal, dorsocentral, and posterior acrostichal areas and lightest on the
humeral, antealar, and supraalar areas and the scutellum; integument of the
pleuron predominantly pale brown, with a variable, irregular darker brown
area on the sternopleuron; patches of pale scales on the upper mesepimeron
and upper and posterior sternopleuron. Wing. All dorsal wing scales dark
brown. Legs. Anterior surface of the hind femur predominantly white,
with a broad, subapical dark band and an apical tuft of white scales; hind tibia
basically dark with a narrow basal white band and occasionally with a median
stripe of pale scales and a narrow apical pale band; hind tarsus dark brown,
with a narrow basal pale band on tarsomeres I-III; mid femur dark, witha
234 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C.(Culex ) whitei
Bram: Genus Culex in Thailand 235
narrow pale basal band and a narrow apical tuft of pale scales; mid tibia and
tarsus similar in color to the hind legs but with the pale scales more sparse;
fore femur similar to the mid femur; fore tibia dark but with an apical tuft of
several pale scales; fore tarsus marked as the hind legs. Abdomen. Terga
dark scaled with moderately broad convex pale basal bands on segments II-
Vill and with basolateral triangular white patches not visible from above;
sterna variable, predominantly pale.
MALE. Head. Proboscis with a small basomedian tuft of short setae;
palpus dark, with a narrow apical band of pale scales on Segment II, a narrow
basal and broad median band of pale scales on III, a narrow basal band of
pale scales on IV, and narrow basal and apical pale bands on V. Terminalia.
(Figure 88C). Subapical lobe of the basimere well developed, with 3 sub-
equal, hooked rods followed by 4 accessory setae (one of which is very broad),
a broad, striated leaf, and a gently curved seta; distimere somewhat ex-
panded, with minute annulations on the convex surface of the apex; inner
division of the phallosome with 4 or more strong, pointed teeth, without a
spiculate portion; outer division of the phallosome with 3 or more strong
teeth similar to those of the inner division; proctiger crowned with a strong
tuft of spines, and with 2 or 3 cercal setae present; basal sternal process
well developed and strongly curved.
LARVA. (Figure 88A,B). Head. Antenna with a narrow, dark basal
band and slightly darker beyond insertion of hair 1-A; head hair 1-C strongly
pigmented, robust, gradually tapering to a sharp point, its length approximate-
ly half the distance between the bases of the pair; 4-C single, simple; 5, 6-C
with 3 or 4 pectinate branches. Thorax. Integument glabrous; hairs 1,2, 3-P
single, pectinate, subequal in length; 4-P bifid, pectinate; 5,6-P single,
pectinate; 7-P usually with 3 branches, but with up to 5 branches, pectinate;
8-P bifid, pectinate; 14-P single, simple. Abdomen. Integument glabrous;
comb variable, consisting of approximately 40 scales arranged in a broadly
triangular patch; individual comb scale elongated, with a prominent apical
Spine and fringed laterally with spicules which become progressively smaller
basally; siphon index variable, ranging from 5:1 to 7:1; 4 or 5 pairs of sub-
ventral tufts inserted on the siphon, the subapical tuft inserted laterally out
of line; individual tufts usually bifid, always simple, their length equal to or
less than the width of the siphon at the point of insertion; pecten consisting
of from 8 to 12 teeth restricted to the basal fourth or less of the siphon; in-
dividual pecten tooth with a prominent apical spine and from 5 to 8 strong
lateral barbs.
TYPE DATA. Lectotype hereby designated: syntype female, "India,
Haflang Hill, Assam, VIII, 1922, Capt. P. J. Barraud, BM 1923-523", in
the British Museum.
DISTRIBUTION. In THAILAND, whitei has been collected from:
Chiang Mai, Chiang Rai, Lampang, Mae Hong Son, Nakhon Ratchasima,
Phet Buri, and Ranong. This species has also been recorded from INDIA
and NEPAL. Specimens have been examined in the U. S. National Museum
—_— eee
Figure 88. C. (Culex) whitei. A, dorsoventral aspect of the head and
prothorax of the fourth stage larva; B, lateral aspect of the
— terminal abdominal segments of the fourth stage larva; C,
dorsal aspect of the male terminalia.
236 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
collection from Ba Long and An Khe, SOUTH VIETNAM**.
During this study the following material has been examined from
Thailand: 4 females and 1 male with their associated larval and pupal skins,
and 32 larvae.
TAXONOMIC DISCUSSION. This species is most easily identified in
the larval stage and may be separated from its closest relative, alienus, by
the short subventral tufts of the siphon and by head hairs 5, 6-C having 3 or 4
branches.
BIOLOGY. Virtually nothing is known of the biology of this apparently
rare species. The larval habitat appears to be limited to ground water ac-
cumulations. Host preferences of the adults are unknown.
SITIENS SUBGROUP
ADULT. Indistinguishable from the vishnui subgroup.
LARVA. Head. Hair 1-C broad, somewhat flattened, its apex rounded
or irregular. Abdomen. Anal gills short, bulbous, their length less than that
of the saddle.
LARVAL HABITAT. Larvae are normally found in brackish water,
usually tidal marshes, and are restricted to coastal regions.
TAXONOMIC DISCUSSION. Although distinguishing features have not
been found which could separate the adults of the sifiens subgroup from other
subgroups within the sztiens group, the 2 distinctive anatomical features of the
larva seem sufficiently unique to warrant separate consideration. The form
of head hair 1-C as noted above is not known in any other members of the sub-
genus; the bulbous anal gills are also unique to the subgroup and probably
represent adaptations to the brackish larval habitat.
Five species fall naturally within this subgroup: sz/zens Wiedemann,
roseni Belkin, whittingtoni Belkin, litovalis Bohart, and neolitoralis, n. sp. ;
however, only 2 of these species are known to occur in Thailand, namely
sitiens and neolitoralis. Belkin (1962) recognized the 4 previously known
species as the szfzens complex of the sifzens subgroup of the szizens group
and included as other members of the sitiens subgroup annulirostris Skuse
and omani Belkin (neither of which are known to occur in Thailand).
CULEX (CULEX) NEOLITORALIS, N. SP.
(Figure 89)
Culex (Culex) litoralis of Colless 1957, Med. J. Malaya 12: 465 (L*).
Although this species is clearly a member of the szfzens subgroup, the
adult female is indistinguishable from alienus. The adult male may be
Figure 89. C. (Culex) neolitoralis. A, dorsoventral aspect of the head
and prothorax of the fourth stage larva; B, lateral aspect of
the terminal abdominal segments of the fourth stage larva; C,
dorsal aspect of the male terminalia with enlarged insert of
the lateral aspect of the inner division of the phallosome.
je
237
Bram: Genus Culex in Thailand
Fig.89
—
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C. (Culex ) neolitoralis
238 Condy. Amin Bint Meets wel yok As A967
recognized by the distinctive inner division of the phallosome of the terminalia.
The fourth stage larva is distinguished on the basis of the unusual structure
of the anal saddle and the pectinate subventral tufts of the siphon.
FEMALE. Head. Proboscis dark brown to black, with a moderately
broad median band of pale scales; palpus similar in color to the proboscis,
but without pale scales; decumbent scales of the vertex pale golden brown at
the occiput, becoming lighter at the orbital line; erect scales golden brown at
the occiput, but dark brown to black posteriolaterally. Thorax. Scutum
basically dark brown but with variable groupings of golden scales which are
particularly dense at the scutal angle, the posterior dorsocentral area, and
the supraalar area, very sparse in the prescutellar space; scutellum with
golden scales prominent on the midlobe; integument of the pleuron usually
pale brown with variable, indistinct darker patterns on the posterior spiracular
plate, the posterior sternopleuron, and the anterior mesepimeron; patches of
pale scales present on the upper sternopleuron, posterior sternopleuron,
anterior meSepimeron, and upper mesepimeron. Wing. All dorsal wing
scales dark brown. Legs. Anterior surface of the hind femur mainly pale
but with a broad, very dark subapical band which extends proximally to form
a dark stripe along the dorsal border; hind tibia dark with a narrow apical
band of pale scales; hind tarsus dark, with narrow pale basal bands on
tarsomeres I-IV; fore and mid femora dark, appearing somewhat paler in the
median area, and with a small apical patch of pale scales; fore and mid tibiae
and tarsi marked as the hind legs. Abdomen. Terga dark, with narrow,
slightly convex pale basal bands on segments III- VII and with a median spot
on segment II; with rather broad basolateral triangular patches which are not
visible from above; sterna dark with broad, pale basal bands.
MALE. Head. Proboscis with a basomedian tuft of a few short setae;
palpus dark with a narrow apical pale band on segment II, narrow basal and
broad median pale bands on III, a narrow basal pale band on IV, and narrow
basal and apical pale bands on V. Terminalia. (Figure 89C). Basimere with
the subapical lobe well developed, with a blunt basal rod and 2 hooked rods
followed by 3 hooked accessory setae, a rather narrow leaf, and a gently
curved seta; distimere normal in shape, without distal annulations; lateral
plate of the phallosome complex, with 5 straight, strong teeth, the spiculate
portion developed away from the teeth, with a prominent projection exhibiting
fine denticles; proctiger crowned with a strong tuft of spines and with 3 cercal
setae; basal sternal process strong, but rather short and straight.
LARVA. (Figure 89A,B). Head, Antenna with a narrow, dark basal
ring and considerably darker beyond insertion of hair 1-A; head hair 1-C
variable in shape, but usually broad, rather bluntly rounded, and somewhat
flattened, its length slightly greater than half the distance between the bases
of the pair; 4-C single, simple; 5-C with from 3 to 5 branches, pectinate; 6-C
with 3 or 4 pectinate branches. Thorax. Integument glabrous; hairs 1,2,3-P
Single, finely pectinate, subequal in length; 4-P bifid, finely pectinate; 5,6-P
single, pectinate; 7-P trifid, pectinate; 8-P bifid, simple; 14-P single,
simple. Abdomen. Integument glabrous; comb consisting of from 20 to 35
elongate, fan-shaped scales arranged in a triangular patch; siphon index
variable, ranging from 2.5:1 to 3.5:1 (average, 3:1); from 6 to 8 pairs of
subventral tufts inserted on the siphon; the apical and antepenultimate tufts
smaller than the other tufts, with from 1 to 3 simple branches, and inserted
laterally out of line; the other tufts with from 4 to 7 branches, pectinate, their
Bram: Genus Culex in Thailand 239
length greater than the width of the siphon at the point of insertion, and inserted
almost in a line so that superficially they do not appear as pairs; pecten consist-
ing of 7 to 13 teeth restricted to the basal third or less of the siphon; individual
pecten tooth with a prominent apical spine and from 2 to 7 coarse lateral barbs;
saddle incomplete, consisting of 2 sclerotized plates, the dorsal plate normal,
the ventral plate triangular, frequently meeting the dorsal plate at the proximal
margin; anal gills bulbous, shorter than the saddle.
TYPE DATA. Holotype male (associated larval and pupal skins and
terminalia slide-mounted) from Ban Laem Sing, Chanthaburi, Thailand, 6
November 1965, from brackish rock pools on a beach, coll. Somboon
Maneechai, deposited in the U. S. National Museum, No. 69351. Paratypes:
3 females and 2 males with their associated pupal skins slide-mounted, and
3 larvae, all with the same data as the holotype.
DISTRIBUTION. In addition to Chanthaburi, the type locality, speci-
mens have also been collected in THAILAND from Ko Si Chang, Chon Buri.
Specimens in the U. S. National Museum collection have been examined
from Finschafen, NEW GUINEA. Colless (1957d) reported what is un-
questionably this species from SINGAPORE harbor, and reported examining
specimens from Seroei, DUTCH NEW GUINEA. Adults and larvae of this
species have been studied from Con Son, SOUTH VIETNAM.
During this study the following specimens were examined in addition to
the type series: 15 females, 13 males, and 54 larvae.
TAXONOMIC DISCUSSION. Colless (1957d) recognized the exceptional
saddle on specimens collected from Singapore, but concluded that this was a
normal feature of /ztovalis Bohart. Although the male terminalia of
neolitoralis and litoralis are strikingly similar, distinctive differences exist
in the adult female and fourth stage larva. The unusual larval saddle of
neolitoralis contrasts the normally incomplete saddle found in litovalis. In
the adult, colorational patterns of the occiput, scutum, abdomen, and legs
differ significantly between the 2 species. Another closely related species is
whittingtoni Belkin, a species known only from the larval stage; differences
exist, however, in the siphon index, chaetotaxy of the head and prothorax,
and in the shape of the saddle.
BIOLOGY. Larvae and pupae of this species were collected from the
type locality during the month of November. Eight of these collections were |
from brackish rock pools near a salt marsh, 1 collection was from a depres-
sion in the salt marsh itself, and 1 collection was made from a large tree hole
within the salt marsh. The material from Chon Buri was collected from
brackish rock pools. Colless (1957d) collected material at Singapore during
June to October, but did not specify the habitat. The specimens from New
Guinea were collected during December from coral pools. Nothing is known
of the abult habits.
CULEX (CULEX) SITIENS WIEDEMANN 1828
(Figures 90, 91, and 92)
Culex sittens Wiedemann 1828, Aussereurop. zweifl. Ins. 1: 542 (?); Theo-
bald 1901, Mon. Cul. 1: 360 (@*); Giles 1902, Handb., 2nd ed.: 400
(?); Blanchard 1905, Les Moustiques: 293 (¢); Leicester 1908, Cul.
Malaya: 143 (co, ?); Theobald 1910, Mon. Cul. 5: 324 (key); Edwards
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
240
C.(Culex) sitiens
Bram: Genus Culex in Thailand 241
1913, Bull. ent. Res. 4: 232 (taxonomy); Barraud 1924, Ind. J. med.
Res. 11: 993 (o*, 2); Barraud 1924, Ind. J. med. Res. 12: 427 (L*);
Buxton and Hopkins 1927, Res. Polynesia and Melanesia 3: 79 (L,
P*); Borel 1930, Monogr. Coll. Soc. Pat. exot. 3: 330 (o'*, 9, L*).
Culex impellens Walker 1859, Proc. Linn. Soc. Lond. 4: 91 (?); Theobald
1901, Mon. Cul. 1: 362 (2*); Giles 1902, Handb., 2nd ed.: 405 (c'*,
°); Theobald 1903, Mon. Cul. 3: 161 (c’); Blanchard 1905, Les
Moustiques: 294 (?); Leicester 1908, Cul. Malaya: 142 (?); Theobald
1910, Mon. Cul. 5: 331 (distribution); Edwards 1913, Bull. ent. Res.
4: 232 (synonymy).
Culex microannulatus Theobald 1901, Mon. Cul. 1: 353 (o*, ?*); Blanchard
1905, Les Moustiques: 292 (?); Leicester 1908, Cul. Malaya: 140
(o,?); Theobald 1910, Mon. Cul. 5: 329 (distribution, taxonomy);
Edwards 1913, Bull. ent. Res. 4: 232 (synonymy).
Culex gnophodes Theobald 1903, Mon. Cul. 3: 163 (2); Leicester 1908, Cul.
Malaya: 145 (?); Edwards 1913, Bull. ent. Res. 4: 232 (synonymy).
Culex somaliensis Neveu- Lemaire 1906, Arch. Parasit., Paris 10: 254
(o*, 9*); Edwards 1911, Bull. ent. Res. 2: 261 (taxonomy); Edwards
1913, Bull. ent. Res. 4: 232 (synonymy).
Culex nigricephala Leicester 1908, Cul. Malaya: 149 (“, 9); Edwards 1932,
in Wytsman, Genera Insect., fasc. 194: 204 (synonymy).
Culex salus Theobald 1908, Rep. Wellcome trop. Res. Lab. 3: 256 (o*, 9*);
Theobald 1910, Mon. Cul. 5: 338 (o'*, 9*, as salsus); Edwards 1913,
Bull. ent. Res. 4: 232 (synonymy).
Culex jepsoni Theobald 1910, Entomologist 43: 158 (¢*); Edwards 1924, Bull.
ent. Res. 14: 394 (synonymy).
Culex saibaii Taylor 1912, Rep. Comm. publ. Hlth. Qd.: 28 (?); Edwards
1924, Bull. ent. Res. 14: 394 (synonymy).
Culicelsa paludis Taylor 1913, Rep. Aust. Inst. trop. Med. 1911: 56 (9);
Taylor 1915, Proc. Linn. Soc. N.S.W. 40: 181 (o); Edwards 1924,
Bull. ent. Res. 14: 394 (synonymy).
Culicada annulata Taylor 1914, Trans. R. ent. Soc. Lond. 1913: 689 (c, ?*);
Edwards 1924, Bull. ent. Res. 14: 394 (synonymy).
Culicelsa annulirostris var. milni Taylor 1914, Trans. R. ent. Soc. Lond.
1914: 196 (o*, 9); Edwards 1924, Bull. ent. Res. 14: 394 (synonymy).
Culex (Culex) sitiens Wiedemann: Edwards 1932, in Wytsman, Genera Insect. ,
fasc. 194: 204 (taxonomy); Barraud 1934, Fauna Brit. India, Diptera
5: 398 (o*, 9, L*); Brug 1934, Bull. ent. Res. 25: 514 (?); Bonne-
Wepster and Brug 1937, Geneesk. Tijdschr. Ned. -Ind. 77: 67 (, 9*);
Bonne-Wepster and Brug 1939, Geneesk. Tijdschr. Ned. -Ind. 79:
1268 (L*); Edwards 1941, Mosq. Ethiop. Reg. 3: 296 (o*, 2); Williams
1943, Hawaii Planters Rec. 47: 215 (?*, L*); Taylor 1943, Serv.
Publ. Sch. Pub. Hlth. trop. Med. Univ. Sydney 4: 99 (c, 2*, L*, P);
Paine 1943, Fiji Dept. Agric. Bull. 22: 12 (?, P*, L*); Bohart 1945,
Figure 90. C. (Culex) sitiens. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax;
a 3 C, dorsal aspect of the female wing; D, anterior surface of
the female legs; E, dorsal aspect of the female scutum and
scutellum; F, dorsal aspect of the female abdomen.
242 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Navmed 580: 80 (?); Bohart and Ingram 1946, Navmed 1055: 18 (o'*,
L*); Carter and Wijesundara 1948, Ceylon J. Sci. 23: 150 (2, L*);
Penn 1949, Pacif. Sci. 3: 80 (P*); Bick 1951, Pacif. Sci. 5: 421
(ecology); Bonne-Wepster 1954, R. trop. Inst. Amst., Spec. Publ.
111: 123 (o*, 9*, L*); Iyengar: and Menon 1955, Bull. ent. Res. 46: 7
(o*, 9, L*); Mattingly and Knight 1956, Bull. Brit. Mus., Ent. 4:
104 (?, L); Bohart 1956, Ins. Micronesia 12: 82 (o*, 9, P, L*); Hsieh
and Liao 1956, Acta ent. Sinica 6: 375 (o*, L*); Belkin 1962, Mosq.
S. Pacif. 1: 205 («*,?, P*, L*); Lien 1962, Pacif. Ins. 4: 635
(ecology); Delfinado 1966, Mem. Amer. ent. Inst. 7: 151 (o*, 2, L*).
Culex (Culex) salinus Baisas 1938, Mon. Bull. Philipp. Hlth. Serv. 18: 204
(o*, 9*, P*, L); Delfinado 1966, Mem. Amer. ent. Inst. 7: 152
(synonymy).
The adult female may be identified by the dark and pale speckling of
the femora and the relatively narrow pale, basal bands on the abdominal
terga. The male terminalia is characterized by the shape and number of
setae on the subapical lobe of the basimere and by the shape of the lateral
plate of the phallosome. The fourth stage larva may be recognized by the
broad, somewhat flattened head hair 1-C, by the bulbous anal gills, and by
the complete anal saddle.
FEMALE. Head. (Figure 90B). Proboscis dark brown with a
moderately broad median band of pale scales and, in Some Specimens, with
a few randomly scattered white scales proximal to the band; palpus similar
in color to the proboscis, usually tipped with several white scales and with
an occasional white scale on segments I-III; decumbent scales of the vertex
pale golden brown, becoming lighter at the orbital line; erect scales dark
brown, or in Some specimens pale brown at the occiput, dark brown
posteriolaterally. Thorax. (Figure 90B, E). Scutum brown, with a variable,
irregular grouping of lighter and darker scales; lighter scales usually found
on the humeral, posterior fossal, anterior and posterior dorsocentral,
supraalar, and prescutellar areas with pale scales also present on the
scutellum; integument of the pleuron uniformly dark brown, with patches of
pale scales on the upper meSepimeron, and the upper and posterior sterno-
pleuron. Wing. (Figure 90C). Dorsal wing scales predominantly dark,
usually with some pale scales on the posterior border of the costa and oc-
casionally with some pale scales on Sc. Legs. (Figure 90D). Anterior
surface of the hind femur predominantly white, but with some dark scales
interspersed among the white, and with dark scales on the dorsal border,
with a broad subapical dark band, and with a small tuft of yellowish scales at
the apex; hind tibia predominantly dark, with an irregular, broad median
stripe of lighter scales and a moderately broad apical pale band; hind tarsus
dark, with basal pale bands on tarsomeres I-IV and a variable number of
pale scales among the dark ones, particularly on tarsomere I; anterior
surface of mid femur uniformly speckled with dark and light scales and with
avery narrow apical band of pale scales; mid tibia and tarsus marked as in
the hind legs; fore femur with an irregular pattern of dark scales with some
light scales interspersed and with a very narrow apical band of pale scales;
fore tarsus marked as in the hind legs. Abdomen. (Figure 90F). Terga
dark brown with variable basal pale bands on segments I-VI; segment I
either dark or with a median pale basal spot; sterna variable, predominantly
Bram: Genus Culex in Thailand 243
white and usually with a complete, rather broad apical dark band.
MALE. Head. (Figure 90A). Proboscis with a rather strong baso-
median tuft of straight setae; palpus dark, with a narrow, variable apical
pale band on segment II, narrow basal and broad median pale bands on III,
a narrow basal pale band on IV, and basal and apical pale bands on V.
Terminalia. (Figure 91C). Basimere with the subapical lobe well developed,
the 3 rods hooked, well developed, followed by 3 or 4 accessory setae, one
of which is longer and broader than the others, a broad leaf, and a gently
curved seta; distimere normal in shape, frequently with extremely minute
annulations at the apex on the convex surface (not illustrated); inner division
of the phallosome with 5 to 7 strong teeth, the spiculate portion appearing as
a broad knob in lateral aspect (figure 91C); proctiger crowned with a strong
tuft of spines and with 3 cercal setae; basal sternal process strong, long,
and prominently curved.
PUPA. (Figure 91A,B).
LARVA. (Figure 92). Head. Antenna with a narrow dark basal band
and progressively darker beyond insertion of hair 1-A; head hair 1-C strongly
pigmented, variable in shape, but usually broad and somewhat flattened,
eradually tapering to a bluntly rounded or irregular apex, its length approx-
imately half the distance between the bases of the pair; 4-C single, simple;
5-C with from 5 to 7 pectinate branches; 6-C usually with 4 branches, pecti-
nate. Thorax. Integument glabrous; hairs 1,2,3-P single, pectinate, sub-
equal in length; 4-P bifid, pectinate; 5,6-P single, pectinate; 7-P trifid,
pectinate; 8-P bifid, pectinate; 14-P single, simple. Abdomen. Integument
glabrous; comb consisting of at least 30 fan-shaped scales arranged in a
broad, triangular patch; siphon index variable, ranging from 3.5:1 to 5. 5:1;
usually 7 or 8 pairs of subventral tufts inserted on the siphon, one of the
tufts inserted laterally out of line; individual tufts with from 2 to 5 finely
pectinate branches, the apical and laterally placed tufts simple, the others
pectinate, their length greater than the width of the siphon at the point of
insertion; pecten consisting of at least 10 or more teeth restricted to the
basal 1/4 to 1/3 of the siphon; individual pecten tooth with a prominent apical
spine and approximately 9 rather fine lateral barbs; saddle completely
ringing the tenth segment at the base, but deeply indented caudoventrally;
anal gills bulbous, shorter than the saddle.
TYPE DATA. Holotype female of sitiens from Sumatra in the
Zoologisk Museum, Copenhagen, Denmark. Holotype female (in extremely
poor condition) of impellens from Makessar, Celebes in the British Museum.
Lectotype of microannulatus hereby designated: syntype male (terminalia
slide-mounted), ''8.2 1900, Quilon, Travencore, S. India, S. P. James", in
the British Museum. Holotype female of gnophodes from Bruas, Dindings,
Malaya in the British Museum. The type locality of somaliensis is
Djibouti, French Somaliland but the type specimen is probably non-existent.
The type locality of nigricephala is Batu Gajah, (Perak), Malaya but the type
specimen is non-existent. Holotype female (in very poor condition) of salus
from Port Sudan, Sudan in the British Museum. Holotype female of jepsoni
from Suva, Fiji Islands in the British Museum. Holotype female of sazbazt
from Saibai Island, New Guinea in the School of Public Health and Tropical
Medicine, University of Sydney, Australia. Holotype female of paludis from
Townsville, Australia in the School of Public Health and Tropical Medicine,
University of Sydney, Australia. Syntypes of annulaia from Townsville,
244 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.91
oll 7
' ant
\ Mecathecset “ ee oh AD
\ Wie hy s oe Seka pity oS
Gun VRAD, yD) i) is by p nal h rng ve ”
Wh we ! peste f ba a SA Wy ie
SENN Ht \ S47,
ny d . he, a oP a
: 2
C. (Culex) sitiens
Bram: Genus Culex in Thailand 245
Queensland, Australia in the School of Public Health and Tropical Medicine,
University of Sydney, Australia. Syntype male and female of annulirostris
var. milni from Milne Bay, Papua, New Guinea in the School of Public Health
and Tropical Medicine, University of Sydney, Australia.
DISTRIBUTION. In THAILAND, sitiens has been collected from:
Chanthaburi, Chon Buri, Krung Thep, Pathum Thani, Prachuap Khiri Khan,
Ranong, Rayong, Samut Prakan, and Samut Sakhon. This species is widely
distributed throughout the coastal areas of the ORIENTAL REGION, EAST
AFRICA, MADAGASCAR, northern AUSTRALIA, and the PACIFIC ISLANDS.
The following specimens from Thailand were examined during this
study: 81 females and 26 males, 7 with their associated larval and pupal
skins, and 150 larvae.
TAXONOMIC DISCUSSION. The systematic position of this variable
species was reviewed by Belkin (1962) who concluded that this is a single,
plastic species with many different populations that have been separated and
then mixed at different times in the past.
Since the adult female may, at times, be confused with some members
of the vishnui subgroup, final determinations should be based on the male
terminalia and the fourth stage larva.
BIOLOGY. In Thailand, larvae of sitiens have been collected from
salt marshes and from ground water habitats and some artificial containers
with brackish water. Collections were made throughout the year. Bick
(1951) reported that in New Guinea coral pools constituted the predominant
larval habitat, and Belkin (1962) collected larvae in the South Pacific from
brackish and sometimes fresh water in ground pools, coral holes, canoes,
and artificial containers. Other authors (e.g. Bonne-Wepster and Brug
1937 and 1939) have attributed a somewhat wider variety of habitats to the
larvae. Colless (1957c) has suggested that this is the typical species found
in the tidal marsh. In all cases, however, the species is generally regarded
as being restricted to coastal regions.
The females have not been reported to attack man in the New Hebrides
and the Solomons (Belkin 1962), but Paine (1943) reported sitiens as a
vicious biter in Fiji. Colless (1959b) utilized precipitin tests to determine
that the females feed primarily on birds and pig, but additional records
included cow, dog, and man; females have also been fed on chicken, mouse,
and rabbit in the laboratory.
Little is known of the disease relations of this species. Iyengar (1953)
found early stages of Wuchereria malayi larvae in a wild-caught female but
concluded that the species was not capable of transmitting the infection.
Hodes (1946) reported laboratory transmission of Japanese B encephalitis by
Culex jepsont (= sitiens), but the significance of this experiment is probably
not great.
Figure 91. C. (Culex) sitiens. A,B, dorsoventral aspects of the pupa;
é C, dorsal aspect of the male terminalia with enlarged insert
of the lateral aspect of the inner division of the phallosome.
—6.246 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.92
C.(Culex) sitiens
Bram: Genus Culex in Thailand 247
GELIDUS SUBGROUP
ADULT. Thorax. Scutum covered with a dense, conspicuous pat-
tern of silver-white scales on the anterior two-thirds. Abdomen. Pale mark-
ings restricted mainly or entirely to the bases of the terga.
LARVA. Indistinguishable from the vishnui subgroup.
TAXONOMIC DISCUSSION. Edwards (1932) recognized 4 species as
comprising his gelidus-series and 2 of these, gelidus and whitmorei, are
found in Thailand and are here assigned to the gelidus subgroup of the sitzens
group. The gelidus subgroup is extremely close to the vishnui subgroup and
may be separated only on the basis of the distinctive markings of the scutum
in the adult; however, at the present time it is considered best to segregate
these two species until more positive relationships can be established.
CULEX (CULEX) GELIDUS THEOBALD 1901
(Figures 93, 94, and 95)
Culex gelidus Theobald 1901, Mon. Cul. 2: 20 (?*); Giles 1902, Handb., 2nd
ed. : 421 (?); Theobald 1903, Mon. Cul. 3: 180 (distribution);
Blanchard 1905, Les Moustiques: 316 (2); Leicester 1908, Cul.
Malaya: 147 (“, 2); Barraud 1924, Indian J. med. Res. 11: 990 (o'*);
Barraud 1924, Indian J. med. Res. 12: 428 (L*); Borel 1930, Monogr.
Coll. Soc. Pat. exot. 3: 327 («*, 9, L*); Barraud and Christophers
1931, Rec. Malar. Surv. India 2: 282 (distribution); Senior-White
1934, Bull. ent. Res. 25: 560 (biology).
Culex gelidus var. cuneatus Theobald 1901, Mon. Cul. 2: 22 (?*); Theobald
1910, Mon. Cul. 5: 313 (distribution); Edwards 1932, in Wytsman,
Genera Insect. , fasc. 194: 203 (synonymy).
Leucomyia gelida (Theobald): Theobald 1907, Mon. Cul. 4: 372 (o*,
taxonomy); Theobald 1910, Mon. Cul. 5: 312 (distribution).
Leucomyia gelida var. bipunctata Theobald 1907, Mon. Cul. 4: 374 (c);
Edwards 1932, in Wytsman, Genera Insect., fasc. 194: 203
(synonymy).
Theobaldiomyia gelidus (Theobald): Brunetti 1912, Rec. Indian Mus. 4: 462 .
(taxonomy).
Culex (Culex) gelidus Theobald: Edwards 1932, in Wytsman, Genera Insect. ,
fasc. 194: 203 (taxonomy); Barraud 1934, Fauna Brit. India, Diptera
5: 407 (oc *, 9, L*); Bonne-Wepster and Brug 1937, Geneesk. Tijdschr.
Ned. -Ind. 77: 74 (¢, 9*); Baisas 1938, Mon. Bull. Philipp. Hlth.
Serv. 18: 212 (*, 9*, L*, P*); Bonne-Wepster and Brug 1939,
Geneesk. Tijdschr. Ned. -Ind. 79: 1271 (L*); Mondchadskii 1951,
Moscow zool. Inst. Akad. Nauk SSSR 37: 262 (L*); Bonne-Wepster
1954, Roy. trop. Inst. Amst., spec. publ. 111: 119 (o, 9*, L*);
Delfinado 1966, Mem. Amer. ent. Inst. 7: 144 (c*, 9, L*).
Figure 92. C. (Culex) sitiens. Fourth stage larva: dorsoventral view
of the head, thorax and abdomen, and lateral aspect of the
< terminal abdominal segments.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C.(Culex) gelidus
248
Bram: Genus Culex in Thailand 249
The adult female may be recognized by the presence of a dense covering
of silver-white scales on the anterior 2/3 of the scutum and the absence of
silver scales on the scutellum. The male is distinguished by the scales of the
scutum and by the simple’inner division of the phallosome of the terminalia,
which is covered with fine sculpturings. The fourth stage larva exhibits an
expanded siphon and the subventral tufts are all inserted in a line.
FEMALE. Head. (Figure 93B). Proboscis dark brown, witha
moderately broad median band of paler scales; palpus dark brown, in some
Specimens with several white scales randomly scattered among the dark;
decumbent scales of the vertex silver-white at the occiput, then with a narrow
area of dark brown scales, and with dull brownish white scales posterolaterally;
erect scales silver-white at the occiput and dark brown posterolaterally.
Thorax. (Figure 93B,E). Scutum covered with a dense pattern of silver-white
scales which terminate at approximately the level of the wing base; within the
silver-white scale pattern are 2 or 4 variable, round patches of dark brown
scales; the posterior third of the scutum and the scutellum more sparsely
covered with dark brown scales; integument of the pleuron light brown, with
some faintly darker brown areas on the sternopleuron; distinct but sparse
patches of white scales on the upper and posterior sternopleuron. Wing.
(Figure 93C). All dorsal wing scales light brown. Legs. (Figure 93D).
Anterior surface of the hind femur lightly scaled proximally, with a dark
apical band which extends along the dorsal margin; hind tibia clothed with light
brown scales and with a narrow apical light band; hind tarsus light brown,
with narrow basal white bands on tarsomeres I-V; fore and mid femora darker;
fore and mid tibiae and tarsi marked as the hind legs. Abdomen. (Figure 93F).
Terga dark scaled with moderately broad basal light bands which reach to the
lateral margins and with a median, posteriorly directed 'V" which in some
cases almost reaches the posterior margin of the tergum and with lateral tri-
angular patches which are barely visible from above (Specimens with a straight
posterior margin are occasionally found); sterna primarily pale, but with dark
scales at the anterior and posterior margins.
MALE. Head. (Figure 93A). Proboscis with a well defined pale band
near the middle, without a basomedian patch of conspicuous setae; palpus dark,
with narrow basal and broad median pale bands on segment III, and narrow basal
pale bands on IV and V. Terminalia. (Figure 94C). Subapical lobe of the basi-
_ mere well developed with 3 basal rods followed by 3 accessory setae (one of
which is very broad), a broad symmetrical leaf, and a gently curved seta;
distimere expanded medially with minute annulations on the convex surface at
the apex; inner division of the phallosome short, blunt, slightly curved laterally
and heavily sculptured; outer division evenly rounded, spiculate; proctiger
crowned with a tuft of strong spines and with 2 or 3 cercal setae; basal sternal
process long, darkly pigmented, and strongly curved.
PUPA. (Figure 94A,B).
LARVA. (Figure 95). Head, Antenna pale, with a narrow, dark
Figure 93. C. (Culex) gelidus. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax;
face C, dorsal aspect of the female wing; D, anterior surface of
the female legs; E, dorsal aspect of the female scutum and
scutellum; F, dorsal aspect of the female abdomen.
250 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.94
A
C.(Culex ) gelidus
Bram: Genus Culex in Thailand 251
basal ring; head hair 1-C darkly pigmented, tapering to a sharp point, its
length approximately half the distance between the bases of the pair; 4-C
single, simple; 5,6-C trifid, pectinate. Thorax. Integument glabrous; hairs
1,2,3-P single, pectinate, subequal in length; 4-P bifid, pectinate; 5,6-P
single, pectinate; 7-P trifid, pectinate; 8-P bifid, pectinate; 14-P single,
simple. Abdomen. Integument glabrous; comb consisting of from 30 to 40
fan-shaped scales arranged in a broad, triangular patch; siphon index variable,
ranging from 3:1 to 3.5:1, the siphon expanded medially; 4 pairs of subventral
tufts inserted in a straight line on the siphon; individual tufts with from 3 to 6
branches, their length less than the width of the siphon at the point of insertion;
pecten consisting of from 7 to 12 teeth restricted to the basal third of the siphon;
individual pecten tooth elongated, with a sharp apical spine and from 4 to 7
sharp lateral barbs.
TYPE DATA. In the British Museum are 2 female specimens labeled
in Theobald's hand as "type" of gelidus: one from 'Malay Penin. Selangor,
Kelang, A. L. Butler"; the other from "29.11. & 21 12 99, Strait Settlements,
Perak, Taiping, L. Wray Junr.". The first of these is hereby designated
lectotype for Culex gelidus; the latter is hereby designated lectotype for Culex
gelidus var. cuneatus. This decision is based on Theobald's (1901) statement
that the original description of gelidus was prepared from a single female, and
by the fact that his description applies perfectly to the designated lectotype.
Theobald's (1901) description (including locality data) of var. cuneatus conforms
precisely to the above lectotype designated for this variety. Holotype male of
Leucomyia gelida var. bipunctata from "India" in the British Museum.
DISTRIBUTION. This species is probably distributed throughout
THAILAND and during this study specimens have been examined from: Chiang
Mai, Chiang Rai, Chon Buri, Ayutthaya, Krung Thep, Lampang, Lamphun,
Narathiwat, Nonthaburi, Pathum Thani, Phet Buri, Ranong, Rayong, Thon
Buri, Trang, and Udon Thani. In addition, gelidus has been recorded from:
MALAYA, INDONESIA, NEW GUINEA, PHILIPPINES, TAIWAN (?), JAPAN,
CHINA, INDOCHINA, BURMA, NEPAL, INDIA, PAKISTAN, and CEYLON.
During this study the following specimens have been studied from
Thailand: 486 females and 362 males, 2 with their associated larval and pupal
skins, and 154 larvae. .
TAXONOMIC DISCUSSION. Although the female of gelidus resembles
whitmorei, the male terminalia and fourth stage larva are distinctive from —
other members of the subgenus. In gelidus females the silver-white scales of
the scutum terminate at about the level of the wing base, whereas in whitmorei
these scales continue posteriorly through the prescutellar space and onto the
mid-lobe of the scutellum.
BIOLOGY. Larvae of gelidus have been collected from a variety of
habitats associated with temporary or semipermanent fresh ground water
such as pools, puddles, small streams, etc., and have occasionally been
collected from artificial containers such as barrels and water tanks. Colless
(1957c) reported a high level of abundance in hyacinth ponds, but also col-
lected larvae from transient pools with emergent vegetation.
Adult females are vicious biters but feed on man only in the absence of
Figure 94. C. (Culex) gelidus. A,B, dorsoventral aspects of the pupa;
< C, dorsal aspect of the male terminalia.
252 Conitih, Aineko Wel ed. velo. wa ie 1407
Fig.95
12
10
MP
10
9
9
KE Ep 4
——z_m®
PE.
eo
C. (Culex) gelidus
> OO eae ee A Pe Sng eee ee =
Bram: Genus Culex in Thailand 253
other suitable hosts. Colless (1959b) concluded that the adults feed primarily
on the larger domestic animals but show a general lack of interest in man
(Colless 1959c). Scanlon and Esah (1965) reported collecting gelidus females
biting man at 2,500 to 4,500 feet of elevation on a mountain in Chiang Mai.
Japanese B encephalitis virus has been isolated from gelidus females
in Malaya (Gould e¢ al. 1962) and in Thailand (Anon. 1964), and the species is
recognized to be at least a potential vector.
CULEX (CULEX) WHITMOREI (GILES) 1904
(Figures 96 and 97)
Taeniorhynchus whitmorei Giles 1904, J. trop. Med. 7: 367 (9).
Taeniorhynchus argenteus Ludlow 1905, Canad. Ent. 37: 98 (?); Theobald
1910, Mon. Cul. 5: 426 (¢*); Edwards 1913, Bull. ent. Res. 4: 232
(synonymy).
Leucomyia plegepennis Theobald 1907, Mon. Cul. 4: 375 (?); Edwards 1913,
Bull. ent. Res. 4: 232 (synonymy).
Culex albus Leicester 1908, Cul. Malaya: 148 (?); Edwards 1913, Bull. ent.
Res. 4: 232 (synonymy).
Culex loricatus Leicester 1908, Cul. Malaya: 151 (?); Edwards 1917, Bull.
ent. Res. 7: 225 (synonymy).
Culex whitmorei (Giles): Edwards 1913, Bull. ent. Res. 4: 232 (taxonomy);
Edwards 1917, Bull. ent. Res. 7: 225 (taxonomy); Barraud 1923,
Indian J. med. Res. 10: 941 (L*); Barraud 1924, Indian J. med.
Res. 11: 988 (c*, 2).
Culex (Culex) whitmorei (Giles): Edwards 1932, in Wytsman, Genera
Insect., fasc. 194: 203 (taxonomy); Barraud 1934, Fauna Brit.
India, Diptera 5: 406 (o'*, °, L*); Stackelberg 1937, Faune de 1'URSS,
Ins. Dipt. 3: 232 (o*, 2); Bonne-Wepster and Brug 1937, Geneesk.
Tijdschr. Ned.-Ind. 77: 77 (o, 2?*); Baisas 1938, Mon. Bull. Philipp.
Hlth. Serv. 18: 211 (o*, ¢, P*, L*); Bonne-Wepster and Brug 1939,
Geneesk. Tijdschr. Ned.-Ind. 79: 1273 (L*); Bohart 1945, Navmed
580: 81 (?, L); LaCasse and Yamaguti 1950, Mosq. Fauna Japan and
Korea: 211 (o*, 9*, L*, P*); Monchadskii 1951, Moscow zool. Inst.
Akad. Nauk SSSR 37: 281 (L*); Bonne-Wepster 1954, Roy. trop. Inst.
Amst., Spec. Publ. 111: 120 (o, 9*, L*); Hara 1957, Jap. J. exp.
Med. 27: 56 (?*); Delfinado 1966, Mem. Amer. ent. Inst. 7: 155
(o*, 7.41.7).
The adult female may be identified by the prominent silver-white pat-
tern of the scutum which extends through the prescutellar space and onto the
mid-lobe of the scutellum. The adult male is recognized by the above
mentioned characters as well as the distinctive setae on the subapical lobe of
the basimere of the terminalia. The fourth stage larva is characterized by
Figure 95. C. (Culex) gelidus. Fourth stage larva: dorsoventral view of
| the head, thorax and abdomen, and lateral aspect of the
= terminal abdominal segments.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
254
C. (Culex) whitmorei
Bram: Genus Culex in Thailand 255
the siphon shape and the pectinate subventral tufts of the siphon.
FEMALE. Head. (Figure 96B). Proboscis dark brown with a rather
broad median band of white scales; palpus similar in color to the proboscis,
tipped with several white scales at the apex; decumbent scales of the vertex
silver-white; erect scales similar in color to the decumbent scales, but
usually with some darker brown scales posterolaterally. Thorax. (Figure
96B,E). Scutum with a pattern of silver-white scales (usually not as dense
as found in gelidus) which extend through the prescutellar space and onto the
mid-lobe of the scutum; within the silver-white pattern is a pair of indistinct
small brown patches in the anterior dorsocentral area and occasionally a
single, variable brown patch in the posterior acrostichal area; integument of
the pleuron variable, the sternopleuron usually darker than the mesepimeron;
patches of pale scales on the upper mesSepimeron and upper and posterior
sternopleuron. Wing. (Figure 96C). All dorsal wing scales brown. Legs.
(Figure 96D). Anterior surface of the hind femur with a variable scattering
of pale scales medially; hind tibia predominantly dark, usually with a median
stripe of pale scales; hind tarsus dark, with moderately broad basal white
bands on tarsomeres I-V and occasionally with a paler median band on tarso-
mere I; fore and mid femora with scattered pale scales; mid tibia marked as
the hind tibia; fore tibia completely dark, or occasionally with a narrow white
basal patch of scales; fore and mid tarsi marked as the hind tarsus but without
the median band on tarsomere I. Abdomen. (Figure 96F). Terga dark scaled
with variable pale basal bands on segments II-VI; the basal pale bands usually
greatly expanded medially and usually not reaching the lateral margins; apico-
lateral pale patches usually present, but not visible from above; sterna pre-
dominantly pale.
MALE. Head. (Figure 96A). Proboscis without a basomedian patch
of conspicuous setae; palpus dark, with a narrow apical pale band on segment
II, narrow basal and rather broad median pale bands on III, and narrow basal
pale bands on IV and V. Terminalia. (Figure 97C). Subapical lobe of the
basimere well developed, with an exceptionally strong, hooked basal rod fol-
lowed by 2 shorter, hooked rods, 3 or 4 hooked accessory setae, a broad
leaf, and a straight seta; distimere normal in shape, without annulations on
the convex surface; inner division of the phallosome with 3 strong, laterally
directed teeth; proctiger crowned with a tuft of strong spines and with 3 or 4
cercal setae; basal sternal process relatively short, gently curved.
LARVA. (Figure 97A,B). Head. Antenna with a narrow, dark basal
ring and somewhat darker beyond insertion of hair 1-A; head hair 1-C strongly
pigmented, gradually tapering to a sharp point, its length approximately half
the distance between the bases of the pair; 4-C single, simple; 5-C bifid,
pectinate; 6-C usually bifid, sometimes trifid, pectinate. Thorax. Integu-
ment glabrous; hairs 1,2,3-P single, pectinate, subequal in length; 4-P
usually bifid, pectinate; 5,6-P single, pectinate; 7-P trifid, pectinate; 8-P
usually bifid (Sometimes single), pectinate; 14-P single, simple. Abdomen.
Figure 96. C. (Culex) whitmorei. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax; C,
sili dorsal aspect of the female wing; D, anterior surface of the
female legs; E, dorsal aspect of the female scutum and
scutellum; F, dorsal aspect of the female abdomen.
256 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
C. (Culex ) whitmore/
Bram: Genus Culex in Thailand 257
Integument glabrous; comb consisting of from 4 to 8 scales arranged in an ir-
regular row; individual comb scale with a very strong, extended apical spine
and fringed with minute basolateral spicules; siphon index variable, ranging
from 4:1 to 5.5:1, the siphon gradually tapering to a narrow apex and gently
curved; 7 or 8 pairs of subventral tufts inserted on the siphon, the apical and
one median pair inserted laterally out of line; individual tufts bifid, pectinate,
their length greater than the width of the siphon at the point of insertion; the
tufts which are laterally displaced are bifid, simple, shorter and slenderer
than the other tufts; hair 2 of the siphon exceptionally large, frequently with
accessory hairlets arising from the base; pecten consisting of from 6 to 12
teeth restricted to the basal fourth or less of the siphon; individual pecten
tooth with a prominent apical spine and with approximately 10 fine lateral barbs.
TYPE DATA. Holotype female of whitmorei from Pampanga, Luzon,
Philippines in the British Museum. Holotype female of avgenteus from
Pampanga, Luzon, Philippines in the U. S. National Museum. Holotype fe-
male of plegepennis from Kobe, Honshu, Japan in the British Museum. Holo-
type female (badly damaged with head missing) of albus from Kuala Lumpur,
Malaya in the British Museum. Holotype female of loricatus from Kuala
Lumpur, Malaya in the British Museum.
DISTRIBUTION. In THAILAND, this species has been collected from:
Chiang Mai, Chiang Rai, Chon Buri, Krung Thep, Lampang, Lamphun, and
Nakhon Nayok; however, it is probably more extensively distributed within
the country. In addition, whitmorei has been reported from throughout the
ORIENTAL REGION as well as from NEW GUINEA, CHINA, KOREA, JAPAN,
and Maritime Province, U.S.S.R.
During this study the following material has been examined from Thai-
land: 268 females, 52 males, and 10 larvae.
TAXONOMIC DISCUSSION. As indicated under gelidus, the two species
are Similar as adults but may be easily separated on the basis of the male
terminalia and fourth stage larva.
BIOLOGY. Although distributed over a broad geographical region,
whitmorei is encountered rather infrequently and, as a result, little is known
about its biology and behaviorial patterns. Larvae have been collected from
fresh ground water habitats such as pools, rice paddies, and the margins of
slowly moving streams. Delfinado (1966) reported females entering houses
and biting at night. |
BITAENIORHYNCHUS SUBGROUP
ADULT. Thorax. Scutum usually covered with an indistinct pattern
of golden or whitish scales on at least the anterior half. Abdomen. Terga
with white apical bands, with apicolateral triangular patches and basal bands,
or with several segments completely covered with pale scales.
LARVA. Head. Hair 1-C lightly pigmented, rather long and cylindrical.
Figure 97. C. (Culex) whitmorei. A, dorsoventral aspect of the head and
| pr othorax of the fourth stage larva; B, lateral aspect of the
— terminal abdominal segments of the fourth stage larva; C,
dorsal aspect of the male terminalia.
258 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Abdomen. Comb consisting of a few very long, simple scales; siphon relative-
ly long; pecten inconspicuous, consisting of a few teeth restricted to the basal
1/5 of the siphon or less.
TAXONOMIC DISCUSSION. Edwards (1932) assigned 12 species to his
bitaeniorhynchus series and there appears to be no doubt that this is a valid
and distinctive subdivision of the sifiens group. Although Thurman (1959)
recorded cornutus Edwards as being represented in fauna of Thailand, the
record is regarded as questionable and only 3 species, bifaeniorhynchus,
pseudosinensis, and sinensis, are presently recognized as members of the
bitaeniorhynchus subgroup of Thailand.
CULEX (CULEX) BITAENIORHYNCHUS GILES 1901
(Figures 98, 99, and 100)
Culex bitaeniorhynchus Giles 1901, J. Bombay nat. Hist. Soc. 13: 607
(adult); Edwards 1913, Bull. ent. Res. 4: 231 (taxonomy); Edwards
1921, Bull. ent. Res. 12: 337 (?); Edwards 1922, Indian J. med.
Res. 10: 282 (o*, 2); Barraud 1923, Indian J. med. Res. 10: 936 (L*);
Barraud 1924, Indian J. med. Res. 11: 984 (<*, 2).
Taeniorhynchus ager Giles 1901, Entomologist 34: 196 (9); Giles 1902,
Handb., 2nd ed. : 365 (?); Blanchard 1905, Les Moustiques: 385
(2); Leicester 1908, Cul. Malaya: 168 (¢,°); Edwards 1912, Bull.
ent. Res. 3: 30 (taxonomy); Edwards 1913, Bull. ent. Res. 4: 231
(synonymy).
Culex infula Theobald 1901, Mon. Cul. 1: 370 (?*); Giles 1902, Handb. , 2nd
ed. : 407 (?); Blanchard 1905, Les Moustiques: 298 (¢); Leicester
1908, Cul. Malaya: 146 (?); Edwards 1912, Bull. ent. Res. 3: 30
(synonymy with ager).
Taeniorhynchus tenax Theobald 1901, Mon. Cul. 2: 198 (?*); Giles 1902, ©
Handb. , 2nd ed. : 365 (?); Theobald 1903, Mon. Cul. 3: 258 (9*);
Blanchard 1905, Les Moustiques: 386 (¢); Leicester 1908, Cul.
Malaya: 167 (?); Edwards 1913, Bull. ent. Res. 4: 231 (synonymy);
Edwards 1922, Indian J. med. Res. 10: 283 (o*,, as variety of
bitaeniorhynchus) ; Barraud 1924, Indian J. med. Res. 11: 985 (o*, 9);
Barraud 1934, Fauna Brit. India, Diptera 5: 393 (?, L); Bohart 1956,
Ins. Micronesia 12: 84 (synonymy).
Grabhamia ambiguus Theobald 1903, Mon. Cul. 3: 248 (o); Edwards 1913,
Bull. ent. Res. 4: 231 (synonymy); Edwards 1922, Indian J. med.
Res. 10: 283 (2, as variety of bitaeniorhynchus); Barraud 1924,
Indian J. med. Res. 11: 985 (?, L); Barraud 1934, Fauna Brit.
India, Diptera 5: 393 (?); Baisas 1938, Mon. Bull. Philipp. Hlth.
Serv. 18: 211 (); Bohart 1956, Ins. Micronesia 12: 84 (synonymy).
Grabhamia taeniarostris Theobald 1907, Mon. Cul. 4: 299 (¢); Edwards
Figure 98. C. (Culex) bitaeniorhynchus. Adult habitus: A, lateral
aspect of the male head; B, lateral aspect of the female head
and thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
ae
Bram: Genus Culex in Thailand 259
C. (Culex) bitaeniorhynchus
260 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
1913, Bull. ent. Res. 4: 231 (synonymy).
Taeniorhynchus tenax var. ocellata Theobald 1907, Mon. Cul. 4: 488 (?*);
Edwards 1913, Bull. ent. Res. 4: 231 (synonymy).
Oculeomyia sarawaki Theobald 1907, Mon. Cul. 4: 515 (¢*); Edwards 1913,
Bull. ent. Res. 4: 231 (synonymy).
Taeniorhynchus domesticus Leicester 1908, Cul. Malaya: 169 (<, 9);
Edwards 1913, Bull. ent. Res. 4: 231 (as variety of bitaeniorhynchus);
Barraud 1924, Indian J. med. Res. 11: 984 (synonymy); Edwards
1932, in Wytsman, Genera Insect., fasc. 194: 202 (synonymy with
auvantapex Edwards). NEW SYNONYMY.
Culicelsa abdominalis Taylor 1913, Rep. Aust. Inst. trop. Med. 1911: 7
(2*): Edwards 1913, Bull. ent. Res. 4: 231 (synonymy).
Culex karatsuensis Mochizuki 1913, Fukuoka Acta med. 7: 28 (o'*, 9*, E);
Tokunaga 1944, Iyo konchu gaku 1: 722 (as variety of bitaeniorhynchus);
LaCasse and Yamaguti 1950, Mosq. Fauna Japan and Korea: 201
(synonymy).
Culex (Culex) bitaeniorhynchus Giles: Edwards 1932, in Wytsman, Genera
Insect. , fasc. 194: 202 (taxonomy); Barraud 1934, Fauna Brit. India,
Diptera 5: 391 (o*, 2, L*); Hopkins 1936, Mosq. Ethiop. Reg. 1: 198
(L); Bonne-Wepster and Brug 1937, Geneesk. Tijdschr. Ned. -Ind.
77: 73 (¢,°%); Baisas 1938, Mon. Bull. Philipp. Hlth. Serv. 18: 211
(o*, 9*, P*, L*); Edwards 1941, Mosq. Ethiop. Reg. 3: 290 (<*, 2);
Bohart 1945, Navmed 580: 78 (¢, L); Bohart and Ingram 1946, Navmed
1055: 77 (o«*, 9, L*); Mohan 1950, Indian J. Malar. 4: 167 (?, P*, L,
E); LaCasse and Yamaguti 1950, Mosq. Fauna Japan and Korea: 201
(o*, 9*, L*, P*); Monchadskii 1951, Moscow zool. Inst. Akad. Nauk
SSSR 37: 258 (L*); Asanuma and Nakagawa, 1953, Misc. Rep. Res.
Inst. nat. Resour. 31: 93 (P*); Laird 1954, Bull. ent. Res. 45: 288
(L*); Belkin 1962, Mosq. S. Pacif. 1: 215 (o*, 2, L*, P*); Delfinado
1966, Mem. Amer. ent. Inst. 7: 140 (o*, 9, P, L*).
The adult female, male terminalia, and fourth stage larva of this
species are extremely variable. The female may be recognized by the combi-
nation of apical tergal bands and the scattered white wing scales. The male
terminalia is characterized by the setae on the subapical lobe of the basi-
mere, and the expanded spiculate portion of the lateral plate of the phallo-
some. The fourth stage larva may be recognized by the distinctive mentum,
by the elongated comb scales, and by the reduced pecten.
FEMALE. Head. (Figure 98B). Proboscis dark brown with a broad
median band of white scales, in Some specimens with a few scattered white
scales proximal to the band; palpus similar in color to the proboscis, with
some scattered white scales, particularly at the apex; decumbent scales of
the vertex light brown, somewhat lighter at the orbital line; erect scales
darker brown. Thorax. (Figure 98B,E). Scutum extremely variable, ranging
from the condition in which all scales are dark brown (illustrated in figure
98E) to a condition in which there exists an indistinct grouping of golden scales
which terminates at about the level of the wing base; a variety of intermediate
forms exist and in some specimens indistinct darker and lighter patches may
be recognized; integument of the pleuron basically light brown, but with in-
distinct and variable darker patterns sometimes present, particularly on the
posterior spiracular plate and the sternopleuron; scattered variable light
Bram: Genus Culex in Thailand 261
scale patches on the posterior pronotum, anterior pronotum, sternopleuron,
and propleuron (in figure 98B illustrated on the sternopleuron only). Wing.
(Figure 98C). All dorsal wing scales usually equally mixed with dark brown
and white scales; this condition is extremely variable and in some speci-
mens there may be only a few pale scales present, or in other specimens
concentrated particularly on the costa, subcosta and vein R,; scales of the
plume series usually broadly clavate, but narrow in some specimens. Legs.
(Figure 98D). Hind femur and tibia basically dark brown, usually with nar-
row basal and apical pale bands and with a variable number of pale scales
randomly scattered among the dark; hind tarsus dark, but also with some
randomly scattered pale scales and with narrow basal pale bands on tarso-
meres I-V and with very narrow apical pale bands on tarsimeres I-IV;
markings of the fore and mid legs similar to the hind legs. Abdomen. (Figure
98F). Terga extremely variable, most commonly dark with a distinct, broad
apical pale band on all segments and with some light scales scattered among
the dark; the apical band is extended anteriorly at the lateral margins; in
some specimens (the "tenax' form), the proximal terga are almost entirely
dark and only a few of the distal terga demonstrate pale apicolateral patches,
and in other specimens (the "ambiguus" form) there are median basal as well
as apical lateral triangular pale patches, or occasionally with the distal seg-
ments completely pale; sterna predominantly pale, but with Melua apical
and basal dark patches.
MALE. Head. (Figure 98A). Proboscis banded and with a strong
medioventral tuft of setae; palpus dark, with variable banding, usually with
a basal pale band on segment II, basal and median pale bands on III, a basal
pale band on IV, and basal and apical pale bands on V. Terminalia. (Figure
99C). Subapical lobe of the basimere rather flat, with a broad basal rod, 2
more Slender rods, 3 or 4 accessory setae, one of which is rather broad
and leaf-like, a slender, short leaf, and a gently curved seta; distimere
normal in shape, but rather short and without annulations; inner division of
the phallosome with from 1 to 3 very broad, blunt teeth, and with the spiculate
portion greatly expanded and extending to a point; proctiger crowned with a
tuft of strong spines and with 2 or 3 cercal setae; basal sternal process short,
usually straight.
PUPA. (Figure 99A, B).
LARVA. (Figure 100). Head. Antenna with a narrow, dark basal
ring and progressively darker beyond insertion of hair 1-A; head hair 1-C
lightly pigmented, its length approximately equal to the distance between the
bases of the pair; 4-C usually bifid, but may range from 1 to 4 branches,
simple; 5-C usually trifid, occasionally with from 2 to 5 branches, pectinate;
6-C almost always bifid, occasionally single or trifid, pectinate; mentum
darkly pigmented, triangular in shape, consisting of 40 or more narrow,
bluntly rounded, tightly compacted denticles on each side. Thorax. Integu-
ment glabrous; hairs 1,2,3-P single, pectinate, subequal in length; 4-P
usually bifid, occasionally single, pectinate; 5,6-P single, pectinate; 7-P
trifid, pectinate; 8-P bifid, pectinate; 14-P single, simple. Abdomen. Inte-
gument glabrous; comb consisting of from 4 to 8 scales arrangedin an ir-
regular row; individual comb scale a single, very strong, elongated, sharply
pointed spine with a basolateral series of fine spicules on each side; siphon
index variable, in specimens from Thailand ranging from 5:1 to 9:1; usually
4, occasionally 3 or 5 pairs of subventral tufts inserted in a line on the siphon;
262 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Fig.99
C. (Culex ) bitaeniorhynchus
Bram: Genus Culex in Thailand 263
individual tufts usually bifid, simple, occasionally with from 1 to 4 branches,
their length less than the width of the siphon at the point of insertion; pecten
consisting of from 4 to 10 teeth restricted to the proximal fifth or less of the
siphon; individual pecten tooth with a strong but short apical spine and from
2 to 5 strong lateral barbs.
TYPE DATA. The type locality of bzfaeniorhynchus is Travancore,
India but the type specimen has been lost. Although Giles' (1901) original
description of ager was of a female, there are 2 males in the British Museum
labeled as "'type" and from these 2 specimens a lectotype is hereby designated:
type male (terminalia mounted and attached to pin), "Capt. Cornwall, Madras,
Paddy field, 24.12.99" in the British Museum. Holotype female of zn/fula
from Taipang, Perak, Malaya in the British Museum. Holotype female of
tenax from Perak, Malaya in the British Museum. Holotype male (terminalia
mounted and attached to pin) of ambiguus from Quilon, Travancore, India in
the British Museum. Holotype female of taeniarosiris from Peradeniya,
Ceylon in the British Museum. Holotype female of fenax var. ocellata from
Kuching, Sarawak, Borneo in the British Museum. Holotype female of
savawaki from Sarawak, Borneo in the British Museum. Lectotype of
domesticus hereby designated: cotype male (terminalia slide-mounted), "Fed.
Malay States, G. F. Leicester, 1912-350", in the British Museum. The type
locality of abdominalis is Ayr and Townsville, Queensland, Australia, but
the type specimens are non-existent. The type locality of kavatsuensis is
Karatsu, Kyushu, Japan but the location of the type specimen is unknown.
DISTRIBUTION. This species is widely distributed throughout THAI-
LAND and during this study specimens have been examined from: Ayutthaya,
Chanthaburi, Chiang Mai, Chiang Rai, Chon Buri, Chumphon, Kanchanaburi,
Krung Thep, Lampang, Mae Hong Son, Nakhon Nayok, Nakhon Ratchasima,
Nakhon Sawan, Nan, Nrathiwat, Nonthaburi, Pathum Thani, Prachuap Khiri
Khan, Phare, Ranong, Rat Buri, Sava Buri, Satun, Songkhla, Sukhothat,
Surat Thani, Tak, Thon Buri, Trang, Ubon Ratchathani, Udon Thani, and
Uthai Thani. In addition, bitaeniorhynchus has been reported from throughout
the ORIENTAL REGION, large areas of the ETHIOPIAN REGION (including
MADAGASCAR), AUSTRALIA, NEW GUINEA, some islands of the SOUTH
PACIFIC, and the SOVIET FAR EAST.
During this study the following specimens of bitaeniorhynchus have
been examined from Thailand: 103 females and 165 males, 40 with their as-
sociated larval and pupal skins, and 360 larvae.
TAXONOMIC DISCUSSION. A number of authors, most recently
Belkin (1962), have suggested the possibility that bif/aeniorhynchus represents
a complex of species and that at least some of the presently regarded synonyms
are, in fact, valid species. Examination of over 100 collections from through-
out Thailand and comparison with material from other areas of Southeast Asia
has ted to the conclusion that bifaeniorhynchus is an extremely plastic species.
Concordance of anatomical and colorational variations could not be established
and, at least for populations on mainland Southeast Asia, it seems that specia-
Figure 99. C. (Culex) bitaeniorhynchus. A,B, dorsoventral aspect of the
: pupa; C, dorsal aspect of the male terminalia with enlarged
insert of the lateral aspect of the inner division of the phallo-
some.
Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
264
Fig.100
<—
SZ
2S
C. (Culex) bitaeniorhynchus
Ze
Bram: Genus Culex in Thailand 265
tion has not progressed to the degree where variant forms may be validly
designated as species. Future investigations involving sibling rearings,
ecology, and perhaps cytogenetic studies may well indicate a need for the
reevaluation of this interesting species.
BIOLOGY. The larval habitat of bitaeniorhynchus is restricted to
accumulations of water containing Spzvogyrva, a filamentous green algae
(Senior-White 1926). Mohan (1950) summarized observations from laboratory
rearings and also concluded that larvae were incapable of developing to
maturity in the absence of green algae. It was also observed that oviposition
is usually postponed until induced by the presence of the algae. Similarly,
Colless (1957c) collected larvae of this species exclusively in the presence
of green algae from ponds, obstructed streams, and other similar impound-
ments.
Sasa and Sabin (1950) collected adults in Japan most frequently from
traps baited with chicken, followed in importance by dog, goat, chicken and
rabbit, man, and rabbit alone; however, only 57 percent of the mosquitoes
from the chicken baited trap were engorged, 40 percent from dog, 8 percent
from goat, 97 percent from chicken and rabbit, none from man, and 50 per-
cent from rabbit alone. In Singapore, Colless (1959b) found that the origin
of blood meals in females collected from unbiased sources were exclusively
from birds, but also collected females from traps baited with pig, fowl, and
cow, but not man. In contrast, bifaeniorhynchus has been reported biting
man in Russia (Chagin 1948 as quoted in Colless 1959b) and in New Guinea
(Bonne-Wepster 1956). Williamson and Zain (1937) induced this species to
feed on man in Malaya.
In New Guinea, de Rook (1957) reported that the most numerous of 3
culicine species found naturally infected with mature forms of Wuchereria
bancrofti was aCulex bitaeniorhynchus - like mosquito; however, it was sug-
gested that, although the specimens agreed rather well with the morphological
concept of bitaeniorhynchus, the form involved may actually have represented
a distinct, man-feeding species. Williamson and Zain (1937) reported labo-
ratory infection and sporozoite development of Plasmodium vivax in
bitaeniorhynchus, but Mohan (1950) was unable to verify this work.
CULEX (CULEX) PSEUDOSINENSIS COLLESS 1955
(Figures 101 and 102)
Culex (Culex) pseudosinensis Colless 1955, Ann. trop. Med. Parasit. 49:
311 (o*, 2, L*).
The adult female may be separated from other members of the sub-
group by the absence of pale scales on the wing veins, and by the presence of
rather broad, pale basal abdominal bands and by apicolateral triangular
patches which may converge at the center of the terga. The adult male is
recognized by the above characters as well as the rather broad leaf on the
Figure 100. C. (Culex) bitaeniorhynchus. Fourth stage larva: dorsoventral
view of the head, thorax and abdomen, and lateral aspect of
— the terminal abdominal segments.
20G:. . Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
subapical lobe of the basimere in the terminalia. The fourth stage larva is
characterized by the rather short comb scales and the length of the pecten.
FEMALE. Head. (Figure 101B). Proboscis dark brown with a
moderately broad median band of pale scales; palpus similar in color to the
proboscis, but wihout light scales present; decumbent scales of the vertex
pale golden; erect scales predominantly pale golden, but some of the scales
dark posterolaterally. Thorax. (Figure 101B,E). Scutum covered with nar-
row pale golden to white scales anterior to the level of the wing base, with ir-
regular dark patches in the area of the fossa, and completely dark posterior to
the level of the wing base; scutellum with dark scales only; integument of the
pleuron uniformly pale brown; with patches of pale scales on the upper and
posterior sternopleuron and occasionally with some scattered pale scales on
the mesepimeron. Wing. (Figure 101C). All dorsal wing scales uniformly
bronze brown. Legs. (Figure 101D). Anterior surface of the hind femur
predominantly dark brown, with a narrow basal band of golden scales and with
an indistinct grouping of paler scales on the basoventral margin; hind tibia
dark brown, with a few randomly scattered pale scales and with a narrow
apical pale band; hind tarsus dark brown, with narrow basal and apical pale
bands on tarsomeres I-IV and with a narrow basal band only on tarsomere V;
fore and mid femora marked as the hind femur but with occasional scattered
pale scales; fore and mid tibiae and tarsi marked as the hind legs. Abdomen.
(Figure 101F). Terga basically dark scaled, but on terga II-VI with moderately
broad, pale basal bands and triangular apicolateral spots which may meet at
the center on terga V-VII; sterna predominantly pale, with scattered apical
dark scales. |
MALE. Head. (Figure 101A). Proboscis with the median pale band
broader than in the female, and with some pale scales on the apex proximal
to the band; with a basomedian tuft of strong setae; palpus dark, with narrow
basal and broad median pale bands on segment II, narrow basal pale bands
on IV and V, and a broad apical pale band on V. Terminalia. (Figure 102C).
Subapical lobe of the basimere well developed with 3 rods, 3 accessory
setae, arather broad, short leaf, and a gently curved seta; distimere normal
in shape, without annulations; lateral plate of the phallosome with 2 blunt
teeth originating from a common base, with the spiculate portion very large
and extended to both dorsal and ventral points; proctiger crowned with a tuft
of strong spines and with 2 or 3 cercal setae; basal sternal process moderately
long, gently curved.
LARVA. (Figure 102A,B). Head. Antenna with a narrow dark basal
ring and slightly darker beyond insertion of hair 1-A; head hair 1-C lightly
pigmented, cylindrical, pointed apically, its length slightly greater than the
distance between the bases of the pair; 4-C single or bifid, simple; 5-C with
3 or 4 branches, pectinate; 6-C bifid, pectinate; mentum heavily sclerotized,
triangular in shape, consisting of 40 or more narrow, bluntly rounded,
tightly compacted denticles on each side. Thorax. Integument glabrous; hairs
Figure 101. CC. (Culex) pseudosinensis. Adult habitus: A, lateral aspect
of the male head; B, lateral aspect of the female head and
thorax; C, dorsal aspect of the female wing; D, anterior
surface of the female legs; E, dorsal aspect of the female
scutum and scutellum; F, dorsal aspect of the female abdomen.
ae
267
Fig.101
Genus Culex in Thailand
Bram
inensis
,
C. (Culex) pseudos
268 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
1,2,3-P single, pectinate, subequal in length; 4-P bifid, pectinate; 5,6-P
single, pectinate; 7-P trifid, pectinate; 8-P bifid, pectinate; 14-P single,
simple. Abdomen. Integument glabrous; comb variable, consisting of from
4 to 12 scales usually arranged in 2 irregular rows; individual comb scale
with a strong, prominent apical spine and with 6 or more fine basolateral
spicules on each side; siphon index variable, ranging from 5.5:1 to 7.5:1
(average, 6.5:1); 3 irregular pairs of subventral tufts inserted on the siphon,
the basal tuft frequently not paired; individual tufts inconspicuous, with 2 or
3 branches, their length less than the width of the siphon at the point of
insertion; pecten consisting of from 8 to 12 teeth restricted to the base of
the siphon, but usually extending further than in bitaeniorhynchus; individual
pecten tooth with a prominent apical spine and 1 or 2 strong basolateral barbs.
TYPE DATA. Holotype female from Singapore (Adelphi Estate),
Malaya in the British Museum.
DISTRIBUTION. In THAILAND**, pseudosinensis is known from a
single collection at Waeng, Narathiwat. In addition to the type locality of
SINGAPORE, this species has also been recorded from INDOCHINA and
Specimens have been examined in the U. 8. National Museum collection from
Saigon, SOUTH VIETNAM#**, and Selangor, MALAYA**.
During this study the following specimens have been examined: the
holotype female and a paratype female with their associated larval and pupal
skins, and 2 males, 1 female, and 3 larvae.
TAXONOMIC DISCUSSION. As indicated by Colless (1955) in his
original description, pseudosinensis demonstrates its closest affinities to
bitaeniorhynchus and geminus Colless with superficial similarity to sinensis.
The adult female is almost indistinguishable from both geminus and sinensis
and the fourth stage larva is extremely close to bitaeniorhynchus. The male
terminalia, particularly the spiculate portion of the lateral plate of the phallo-
some, is sufficiently distinctive to permit accurate determination and it is
strongly suggested that final conclusions not be drawn without examination of
this structure. .
BIOLOGY. The type habitat of the larvae was recorded as a moderately
large stream pool just inside the jungle fringe. In Thailand, pupae were
collected from rock pools beside a waterfall. Nothing else is known about the
biology of this apparently rare species. :
CULEX (CULEX) SINENSIS THEOBALD 1903
(Figures 103 and 104)
Culex gelidus var. sinensis Theobald 1903, Mon. Cul. 3: 180 (¢).
Culex sepositus Leicester 1908, Cul. Malaya: 152 (?); Edwards 1913, Bull.
ent. Res. 4: 231 (synonymy).
Taeniorhynchus tenax Leicester 1908 (not Theobald 1901), Cul. Malaya: 167
Figure 102. C. (Culex) pseudosinensis. A, dorsoventral aspect of the
head and prothorax of the fourth stage larva; B, lateral
aspect of the terminal abdominal segments of the fourth
stage larva; C, dorsal aspect of the male terminalia with
enlarged insert of the lateral aspect of the inner division of
the phallosome.
>
Bram: Genus Culex in Thailand 269
C. (Culex) pseudosinensis
270 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
(2); Edwards 1913, Bull. ent. Res. 4: 231 (synonymy).
Leucomyia sinensis (Theobald) 1910, Mon. Cul. 5: 313 (taxonomy).
Culex sinensis Theobald: Edwards 1913, Bull. ent. Res. 4: 231 (taxonomy);
Edwards 1921, Bull. ent. Res. 12: 337 (4,2); Barraud 1924, Indian
J. med. Res. 11: 986 (c*, 2).
Culex tripunctatus Mochizuki 1913, Fukuoka Acta med. 7: 24 (o*, 9*, E);
Edwards 1922, in Wytsman, Genera Insect., fasc. 194: 203 (synonymy).
Culex (Culex) sinensis Theobald: Edwards 1932, 7m Wytsman, Genera Insect. ,
fasc. 194: 203 (taxonomy); Barraud 1934, Fauna Brit. India, Diptera
5: 394 (o*, 2, L); Stackelberg 1937, Fauna de l'URSS, Ins. Diptera 3:
231 (o*); Bonne-Wepster and Brug 1937, Geneesk. Tijdschr. Ned. -
Ind. 77: 78 (%, 2*); Baisas 1938, Mon. Bull. Philipp. Hlth. Serv.
18: 209 (?, E); Bonne-Wepster and Brug 1939, Geneesk. Tijdschr.
Ned. -Ind. 79: 1274 (L*); Bohart and Ingram 1946, Navmed 1055: 79
(**, 9, L*); LaCasse and Yamaguti 1950, Mosq. Fauna Japan and
Korea: 206 (o*, ?*, L*, P*); Monchadskii 1951, Moscow zool. Inst.
Akad. Nauk. SSSR 37: 275 (L*); Bonne-Wepster 1954, Roy. trop.
Inst. Amst. Spec. Pub. 111: 116 (“, 9*, L*); Hara 1957, Jap. J.
exp. Med. 27: 55 (?*); Delfinado 1966, Mem. Amer. ent. Inst. 7:
151.{2).
The adult female may be recognized by the nature of the apical and
basal banding of the abdominal terga and by the uniformly dark dorsal wing
scales. The simple, distinctive phallosome of the male terminalia is
characteristic. The fourth stage larva may be identified by the chaetotaxy of
the head and the shape of the mentum.
FEMALE. Head. (Figure 103B). Proboscis dark brown, witha
rather broad median pale band of scales; palpus similar in color to the
proboscis, tipped with pale scales and occasionally with several randomly
placed pale scales on segments I-III; decumbent scales of the vertex pale
golden, becoming lighter at the orbital line; erect scales golden to pale
golden at the occiput and dark brown posterolaterally. Thorax. (Figure
103B,E). Scutum covered with variable pale brown to white scales anterior
to the supraalar area, dark posterior to the supraalar area with some pale
scales in the prescutellar space, and with variable dark patches on the fossa;
scutellum dark but with some light scales on each of the 3 lobes; integument
of the pleuron light brown, occasionally with variable, indistinct darker areas
particularly on the sternopleuron and posterior spiracular plate; patches of
dull white scales present on the upper meSepimeron and upper and posterior
sternopleuron. Wing. (Figure 103C). Dorsal wing scales usually uniformly
dark brown but with some scattered pale scales and a variable row of white
scales occasionally present on the posterior margin of the costa. Legs.
(Figure 103D). Anterior surface of the hind femur with a narrow pale basal
band, a uniform pattern of light and dark scales, a subapical dark band, and
SS AON ESS REE On TI CE RS OO SERA PAT ETRE DA SS RCE AE SN OE ASR SRE — REST: AREY" SN OY TR SSRN CN ENR WO RE AS ES ENS AD EE ONS Bf ER PEE
Figure 103. CC. (Culex) sinensis. Adult habitus: A, lateral aspect of the
male head; B, lateral aspect of the female head and thorax;
C, dorsal aspect of the female wing; D, anterior surface of
the female legs; E, dorsal aspect of the female scutum and
scutellum; F, dorsal aspect of the female abdomen. :
nes eH Scheid
C. (Culex ) sinensis
Bram: Genus Culex in Thailand
2'T2 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
an apical tuft of pale scales; hind tibia with a variable mixture of dark and
pale scales; hind tarsus dark, with narrow, basal pale bands on tarsomeres
I-IV; markings of the fore and mid legs variable, but similar to those of the
hind legs. Abdomen. (Figure 103E). Terga dark with variable basal and
apical pale bands on segments II-VI; basolateral pale patches present on
each tergum, but not visible from above; sterna predominantly pale, with
mediolateral dark patches.
MALE. Head. (Figure 103A). Proboscis with a small basomedian
tuft of setae; palpus dark, with narrow basal and broad median pale bands on
segment II, with a narrow basal pale band on IV, and narrow basal and apical
pale bands on V. Terminalia. (Figure 104C). Subapical lobe of the basimere
well developed, with 3 hooked rods, 3 fine and 1 strong accessory setae, a
well developed leaf, and a gently curved seta; distimere normal in shape,
without distal annulations; phallosome simple, the inner division gradually
tapering, slightly curved, and spiculate, the arms parallel; outer division
ereatly reduced, represented by a small basal protuberance; proctiger crowned
with a tuft of strong spines and with 2 or 3 cercal setae; basal sternal process
well developed, darkly pigmented, and strongly recurved.
LARVA. (Figure 104A,B). Head. Antenna with a narrow dark basal
ring and progressively darker beyond hairs 2,3-A, which are inserted
considerably below the apex of the shaft; head hair 1-C lightly to moderately
pigmented, its length approximately half the distance between the bases of
the pairs; 4-C single, simple, exceptionally long; 5, 6-C usually bifid, pecti-
nate; mentum triangular, strongly pigmented, with approximately 7 robust,
distinctly separated lateral teeth on each side. Thorax. Integument glabrous;
hairs 1,2,3-P single, pectinate, subequal in length; 4-P very short and fine,
with from 2 to 4 simple branches; 5-P single, pectinate; 6-P single or bifid,
pectinate; 7-P trifid, pectinate; 8-P single, pectinate; 14-P single, simple.
Abdomen. Integument glabrous; comb consisting of from 3 to 5 scales ar-
ranged in an irregular row; individual comb scale a strong, prominent, sharply
pointed spine with a series of fine, basolateral spicules on each side; siphon
index variable, ranging from 6:1 to 8:1; 5 or 6 irregular pairs of subventral
tufts inserted on the siphon; individual tufts with 2 or 3 branches, their length
less than the width of the siphon at the point of insertion; pecten consisting of
from 1 to 5 teeth, restricted to the extreme base of the siphon; individual
pecten tooth with a strong distal spine and from 1 to 5 fine basolateral barbs
on one side, or the basal tooth may consist of a strong distal spine with fine
basolateral barbs on each side.
TYPE DATA. Holotype female of sizensis from Shaohyling, China in
the British Museum. The type locality of sepositus is Kuala Lumpur, Selangor,
Malaya but the type specimen is non-existent. The type locality of tenax is
Kuala Lumpur, Selangor, Malaya but the type specimen is non-existent. The
type locality of tvipunctatus is Fukuoka, Kyushu, Japan but the location of the
type specimen is. unknown.
DISTRIBUTION. In THAILAND, sznvensis has been studied from:
Figure 104. C. (Culex) sinensis. A, dorsoventral aspect of the head and
prothorax of the fourth stage larva; B, lateral aspect of the
terminal abdominal segments of the fourth stage larva; C,
dorsal aspect of the male terminalia. —>
273
Bram: Genus Culex in Thailand
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274 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
Ayutthaya, Chiang Mai, Chon Buri, Narathiwat, Nakhon Nayok, Nakhon Si
Thammarat, Nonthaburi, Prachuap Khiri Khan, Trang, Ubon Ratchathani,
and Udon Thani. The species has also been reported from CHINA, USSR,
KOREA, JAPAN, RYUKYU-RETTO, TAIWAN, PHILIPPINES, INDONESIA,
MALAYA, BURMA, INDIA, and CEYLON. Specimens have been studied in
the U. S. National Museum collection from Qui Nhon, SOUTH VIETNAM**,
During this study the following specimens were examined from Thai-
land: 60 females, 16 males, and 12 larvae.
TAXONOMIC DISCUSSION. Although the adult female demonstrates
a Similarity to pseudosinensis, the male terminalia and fourth stage larva are
clearly distinctive.
BIOLOGY. Larvae are found in a variety of ground water habitats,
frequently in association with green algae; predominant among these are rice
fields, ditches, ponds, and streams. Habits of the adults are poorly known,
but Lien (1962) reported that on Taiwan sinensis enters houses, human baited
net traps, and light traps. Scanlon and Esah (1965) collected this species
biting man up to 4,500 feet near Chiang Mai, Thailand.
MISIDENTIFICATIONS
CULEX (MOCHTHOGENES) CASTRENSIS EDWARDS 1922
Culex castrensis Edwards 1922, Indian J. med. Res. 10: 285 (c*, 9).
Causey (1937) tentatively recorded castrensis from Thailand, but it is
felt that his record actually referred to folzatus, which at that time was
regarded as a subspecies of castrensis.
CULEX (LOPHOCERAOMYIA) FLA VICORNIS BARRAUD 1924
Culex (Lophoceratomyia) flavicornis Barraud 1924, Indian J. med. Res. 12:
45 (o'*, 9).
Specific locality data were not included in the original record of this
Species by Thurman (1959). Specimens in the U. S. National Museum col-
lection from Thailand and determined as /lavicornis by E. Thurman have been
examined and found to represent zncomptus.
CULEX (LOPHOCERAOMYIA) FRAUDATRIX (THEOBALD) 1905
Lophoceratomyia fraudatrix Theobald 1905, Ann. hist.-nat. Mus. hung. 3:
94 (o*, 2*),
Causey (1937) and Iyengar (1953) recorded fraudatrix from Thailand,
however, Colless (1959a) showed that there is no evidence for the occurrence
of this species in the Oriental region. Previous records from Tiailand are
considered to be in error and probably represent macdonaldi and variatus.
Bram: Genus Culex in Thailand 275
CULEX (LOPHOCERAOMYIA) UNIFORMIS (THEOBALD) 1905
Lophoceraomyia uniformis, Theobald 1905, Bombay nat. Hist. Soc. 16: 245
(o*, 9).
Although Thurman (1959) reported that this species had been recorded
from Thailand between 1950 and 1956, no reference can be located which
establishes this record. Specimens in the Thurman collection, determined
as uniformis, have been examined and found to belong to 2 species, minor and
spiculosus. The Thurman record was probably based on these misidentified
specimens.
CULEX (CULICIOMYIA) VIRIDIVENTER GILES 1901
Culex viridiventer Giles 1901, J. Bombay nat. Hist. Soc. 13: 609 (adult).
Thurman (1959) recorded viridiventer from Thailand but did not in-
clude specific locality data. Specimens in the U. S. National Museum col-
lection, determined by E. Thurman as vividiventer, have been examined and
found to represent a new species which is described earlier in this paper as
thurmanorum.
CULEX (CULEX) PIPIENS PIPIENS LINNAEUS 1758
Culex pipiens Linnaeus 1758, Syst. Nat. 10th ed. 1: 602 (adult).
Thurman and Thurman (1955) recorded this subspecies (as pipions in
contrast to quinquefasciatus) from Thailand, but examination of over 800 male
terminalia slides of pipiens sensu lat. (including specimens in the Thurman
collection) from throughout Thailand has not suggested the presence of pipiens
pipiens.
CULEX (CULEX) THEILERI THEOBALD 1903
Culex theileri Theobald 1903, Mon. Cul. 3: 187 (o*, ?*).
Thurman (1959) originally recorded ¢heileri from Thailand but did not
include specific locality data. Specimens in the U. S. National Museum col-
lection from Thailand, determined by E. Thurman as theileri, have been
studied and found to be annulus. No specimens from Thailand have been seen
which can be attributed to this species.
DOUBTFUL RECORDS
CULEX (MOCHTHOGENES) KHAZANI EDWARDS 1922
Culex khazani Edwards 1922, Indian J. med. Res. 10: 286 (c'*).
276 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
The original record of this species for Thailand was that of Thurman
(1959). Although locality data were not included, the material upon which the
report was based was probably collected in northern Thailand. Specimens
determined as this species have not been located in the Thurman collection,
nor have specimens of khazani been seen from any of the Thailand collections.
CULEX (LOPHOCERAOMYIA) MINUTISSIMUS (THEOBALD) 1907
Culiciomyia minutissimus Theobald 1907, Mon. Cul. 4: 235 (9).
As in the case of khazani above, Thurman (1959) recorded minutissimus
from Thailand, but no specimens can be located which can be attributed to
this species.
CULEX (CULEX) CORNUTUS EDWARDS 1922
Culex cornutus Edwards 1922, Indian J. med. Res. 10: 283 (c*, 2).
Thurman (1959) recorded cornutus from Thailand without including
specific locality data. No specimens have been found in the Thurman col-
lection which can be attributed to this species, and no specimens have been
seen from any of the Thailand collections.
CULEX (CULEX) MIMETICUS NOE 1899
Culex mimeticus Noé 1899, Boll. Soc. ent. ital. 31: 240 (¢). —
Causey (1937) recorded mimeticus from the northern hill area of
Thailand. During this study, specimens from Thailand have not been seen
which could be ascribed to this species and it is suggested that Causey's
record may possibly have been a misidentification of mimulus.
CULEX (CULEX) UNIVITTATUS THEOBALD 1901
Culex univittatus Theobald 1901, Mon. Cul. 2: 29 (c*, 9*).
Thurman (1959) recorded univittatus from Thailand without listing
specific locality data. No specimens of this species have been found in the
Thurman collection or in any other material from Thailand.
CULEX (CULEX) VISHNUI THEOBALD 1901
Culex vishnui Theobald 1901, Mon. Cul. 1: 355 (¢*).
Edwards (1922) recorded the distribution of vishnui as throughout the
Oriental region and Barraud and Christophers (1931), Causey (1937), Iyengar
Bram: Genus Culex in Thailand 240
(1953), and Thurman and Thurman (1955) have all recorded the’species from
Thailand. The validity of all references to vishnui prior to 1957 has been
seriously challenged by Colless (1957a) who pointed out that 4 species of the
subgenus, Pseudovishnui, alienus, annulus, and perplexus, were identifiable
in the adult stage as vishnui as defined prior to 1957. Most of the above
records from Thailand probably referred to pseudovishnui or annulus.
ACKNOWLEDGMENTS
Dr. Botha de Meillon, Responsible Investigator, South East Asia
Mosquito Project, and Dr. Alan Stone, Agricultural Research Service, U. S.
Department of Agriculture, both offered many valuable suggestions through-
out this study and critically reviewed and edited the manuscript. To these
men I wish to offer my sincere appreciation and thanks.
Lt. Colonel J. E. Scanlon, Walter Reed Army Institute of Research,
initiated the collections which led to these studies while at the South East
Asia Treaty Organization Medical Research Laboratory, Bangkok, Thailand.
Personnel of the Department of Medical Entomology, SEATO Medical Research
Laboratory collected most of the material and prepared specimens for study.
Included among these were: Dr. D. J. Gould and MSG. E. L. Peyton who
supervised the collection and preparation of specimens and continued their
interest throughout the study; Mr. S. Esah; Mr. S. Maniwongse; Mr. S.
Chunchulcherm; Mr. K. Mongkolpanya; Mr. S. Maneechai; Mr. C. Diraphat;
Mrs. R. Rattanarithikul; Miss S. Sandhinand; and Miss P. Nawarat. Without
the SEATO collection and participation this study would not have been possible.
Technical assistance at SEAMP, Smithsonian Institution, was provided
by Miss C. C. Lipscomb, Miss M. T. Houck, Mr. C. J. Lane, Mr. M.
Rattanarithikul, Miss V. M. Ford, and Mr. A. Uter. Miss T. L. Ford and
Miss J. L. Hwang of the SEAMP prepared a large number of the illustrations.
Illustrations were also prepared by artists of the U. S. Army, 406th General
Medical Laboratory, Tokyo, Japan. Included among these were: Mr. K.
Daishoji, Mr. M. Misaki, Mr. K. Miyasaka, Mr. Y. Morine, and Mr. S.
Ohtawa.
Dr. P. F. Mattingly kindly permitted examination of types and other
specimens in the British Museum (Natural History) and arranged for the loan
of a number of specimens. Dr. D. H. Colless, C.S.I.R.O., Canberra,
Australia assisted in the determination of a number of species in the subgenus
Lophoceraomyia and also loaned several specimens.
To the many individuals who contributed indirectly to the completion
of this study, I wish to express my gratitude.
278 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
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APPENDIX A
SPECIES OF THE GENUS CULEX OCCURRING IN SOUTHEAST ASIA
BUT NOT KNOWN FROM THAILAND!
SPECIES
NEOCULEX
hayashii Yamada
nematoides Dyar & Shannon
okinawae Bohart
simplicicornis Edwards
sumatranus Brug
MOCHTHOGENES
culionicus Delfinado
hackeri Edwards
khazani Edwards
latifoliatus Delfinado
laureli Baisas
tricontus Delfinado
yeagervi Baisas
ACALLEOMYIA
obscurus (Leicester)
DISTRIBUTION
Japan, Ryukyu-Retto,
Taiwan, China, Korea,
USSR.
Philippines.
Ryukyu-Retto.
Borneo.
Sumatra, China.
Philippines.
Malaya.
India, Indochina.
Philippines.
Philippines.
Philippines.
Philippines.
Malaya.
* = stage or sex described and illustrated; X = stage or sex described;
- = Stage or sex unknown; ? = stage or sex described but questionable.
Bram: Genus Culex in Thailand
SPECIES
LOPHOCERAOMYIA
acutipalus Colless
bandoengensis Brug
barkerii (Theobald)
bernardi (Borel)
brevipalpus (Theobald)
coerulescens Edwards
crassicomus Colless
cubitatus Colless
demissus Colless
eminentia (Leicester)
fulleri (Ludlow)
gibbulus Delfinado
hewitti (Edwards)
inculus Colless
jenseni (Meijere)
josephineae Baisas
lavatae Stone & Bohart
minutissimus (Theobald)
navalis Edwards
niger (Leicester)
pachecoi Baisas
DISTRIBUTION
Singapore.
Malaya, Java.
Borneo.
Indochina.
Singapore.
Singapore, Borneo.
Malaya.
Singapore.
Malaya.
Malaya, Singapore,
Borneo.
Philippines.
Philippines.
Borneo, Singapore,
Sumatra.
Malaya.
Singapore, Java,
Sumatra.
_ Philippines.
Philippines.
Ceylon, India, Java,
Borneo, China, Maldive
Is.
Singapore, Borneo.
Malaya, Sumatra.
Philippines.
287
288 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
SPECIES
plantaginis Barraud
tuberis Bohart
uniformis (Theobald)
whartoni Colless
CULICIOMYIA
bahri (Edwards)
javanensis Bonne-Wepster
ryukyensis Bohart
shebbearei Barraud
viridiventer Giles
CULEX
annulirostris Skuse
bihamatus Edwards
diengensis Brug
jJasyi Baisas
geminus Colless
incognitus Baisas
jacksoni Edwards
mimeticus Noé
orientalis Edwards
bropinquus Colless
solitarius Bonne-Wepster
lee]?
“e
~ Mm
I
gees
DISTRIBUTION
India, ? Nepal.
Ryukyu-Retto.
Ceylon, India, Hainan
Is., New Guinea,
Philippines.
Malaya, Singapore.
Ceylon, ? Java.
Java.
Ryukyu-Retto, Japan.
India, Ceylon, Borneo,
China.
India, Pakistan, Nepal.
Indonesia, Philippines,
Australasian region.
Timor.
Java.
Philippines.
Singapore.
Philippines.
China, Korea, Nepal,
USSR.
Oriental region.
Japan, Siberia, Korea,
China, Taiwan.
Singapore.
New Guinea, ? Java.
Since the above appendix was compiled, my attention has been drawn to the
following additional species described from the Andaman Islands: C (Mochthogenes)
shrivastavii Wattal, Kalra, and Krishnan 1966, Bull. Indian Soc. Malar. Com. Dis.
3: 159 (o*),
Bram: Genus Culex in Thailand 289
APPENDIX B
PRESENT STATUS OF THE CULEX FAUNA OF THAILAND!
[fel
SPECIES
LUTZIA
fuscanus Wiedemann
halifaxii Theobald
NEOCULEX
brevipalpis (Giles)
tenuipalpis Barraud
MOCHTHOGENES
foliatus Brug
hinglungensis Chu
malayi (Leicester)
LOPHOCERAOMYIA
aculeatus Colless
alphus Colless
bengalensis Barraud
cinctellus Edwards
curtipalpis (Edwards)
eukrines Bram and
Rattanarithikul
_ fuscosiphonis Bram and
Rattanar ithikul
ganapathi Colless
BIOLOGY
Well known from
Rangoon.
Fairly well known.
Larval habitats known.
Poorly known.
Larval habitats known.
Unknown.
Larval habitats known.
Larval habitats known.
Poorly known.
Poorly known.
Poorly known.
‘Larval habitats known.
Larval habitats known.
Poorly known.
Larval habitats known:
1, = stage or sex described and illustrated; X = stage or sex described;
- = stage or sex unknown; ? = stage or sex described but questionable.
290 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
SPECIES
incomptus Bram and
Rattanarithikul
infantulus Edwards
lucaris Colless
macdonaldi Colless
mammilifer (Leicester)
minor (Leicester)
minor form bicornutus
(Theobald)
peytoni Bram and
Rattanarithikul
pholeter Bram and
Rattanarithikul
quadripalpis (Edwards)
reidi Colless
rubithoracis (Leicester)
spiculosus Bram and
Rattanarithikul
traubi Colless
variatus (Leicester)
wilfredi Colless
THAIOMYIA
dispectus Bram
CULICIOMYIA
bailyi Barraud
barrinus, n. sp.
fragilis Ludlow
a be iw
BIOLOGY
Unknown.
Larval habitats known.
Unknown.
Poorly known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Poorly known.
Poorly known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Bram: Genus Culex in Thailand 291
SPECIES
nigyopunctatus Edwards
pallidothorax Theobald
| papuensis (Taylor)
scanloni, n. Sp.
spathifurca (Edwards)
spiculothorax, n. sp.
termi Thurman
thurmanorum, Nn. sp.
CULEX
alienus Colless
annulus Theobald
barvaudi. Edwards
bitaeniorhynchus Giles
fuscocephala Theobald
gelidus Theobald
hutchinsoni Barraud
mimulus Edwards
neolitoralis, n. sp.
perplexus Leicester
p. quinquefasciatus Say
pseudosinensis Colless
pseudovishnui Colless
2 ath
*| * | * *
BIOLOGY
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats and
some host preferences
known.
Poorly known.
Larval habitats known.
Larval habitats known.
Poorly known.
Fairly well known.
Poorly known.
Fairly well known.
Fairly well known.
Fairly well known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Larval habitats known.
Well known from
Rangoon.
Poorly known.
Larval habitats and
some host preferences
known.
292 Contrib. Amer. Ent. Inst., vol. 2, no. 1, 1967
SPECIES
sinensis Theobald
sitiens Wiedemann
tritaeniorhynchus Giles
whitei Barraud
whitmorei (Giles)
Eloi be al
* | kK | Xx *K
x
x
*
*
*
BIOLOGY
Larval habitats and
some adult biology
known.
Fairly well known.
Fairly well known.
Poorly known.
Poorly known.
293
INDEX
In the following index, all names are listed alphabetically, regardless
of whether they are genus-group or species-group names.
Valid names are
set in roman type, synonyms are italicized. The italicized pages are those
which begin the primary treatment of the species.
Numbers in parentheses
refer to the figures illustrating the species in question.
abdominalis, 260,263
Acalleomyia, 5,8, 286
aculeatus, 47,49,50, 66, 289 (13)
acutipalus, 287
adelae, 205,207
aedes, 39, 42
ager, 258,263
albopleura, 137,141
albus, 253,257
alienus, 6,173,177, 182,202, 236,
277,291 (75)
alphus, 47,49, 53,289 (14)
ambiguus, 258,263
annulata (Culicada), 241,243
annulata (Culiciomyia), 133,137
annulata (Trichopronomyia), 163
annulirostris, 6, 236, 288
annuloabdominalis, 137,141
annulus, 173,177,182, 201, 205, 232,
275,277,291 (76, 77, 78)
Aporoculex, 163
argenteus, 253,257
aureopunctis, 17,22
bahri, 6,288
bailyi, 6,120,122, 123,127,290 (38,
39, 40)
balabacensis (Anopheles), 127
bandoengensis, 82,287
barkerii, 42
barkerii, 287
barraudi, 171,177,181, 209, 291 (79)
barrinus, 6,120, 121,123, 125, 290
(41)
bengalensis, 5,48, 82,95, 289 (24)
bernardi, 287
— bicornuta, 96
bicornutus, 46, 96, 99, 290 (29A')
bigoti, 11
bihamatus, 288
bipunctata, 247,251
biroi, 225, 230
bitaeniorhynchus subgroup, 3, 6, 169,
201, 257
bitaeniorhynchus, 6, 167,169, 175,
179, 258, 265, 268, 291 (98, 99,
100)
brevipalpis, 5,24, 25,29, 31, 289 (6,
7,8)
brevipalpus subgroup, 5,8, 9,11, 44,
iE
brevipalpus, 76,287
Cacoculex, 165
castrensis, 34,317,274
ceylonica, 129,131
chiungchungensis, 93,95
chiyutoi, 34,37
cinctellus, 46, 49,50, 55,289 (15)
cingulatus, 192,196
Coelomomyces, 9
coerulescens, 287
concolor, 11,13,15
confusus, 215,217
cornutus, 258,276
crassicomus, 287
cubitatus, 50, 287
Culex (genus), 2,7
Culex (subgenus), 3, 6,8,9,10, 12, 763,
288,291
Culiciomyia, 3,5, 8,10,11,12, 24, 42,
44,113,117, 288, 290
culionicus, 286
cuneatus, 247,251
curtipalpis, 5, 44, 47, 76,289 (22)
Cyathomyia, 42
debilis, 165
demissus, 82, 90,287
diengensis, 288
dispectus, 8,10,11, 113,290 (35, 36,
37)
doleschallii, 192
dolosa, 163
domesticus, 260, 263
294 Index to Culex of Thailand
edwardsi, 211,213
€leuthera, 165
eminentia, 76,81, 287
eukrines, 45, 48, 85,289 (25)
Eumelanomyia, 23
fasyi, 288
fatigans, 182,192,194, 196
fidelis, 25,29
flavicornis, 274
flavus (Armigeres), 117
foliatus, 5, 34,37,274, 289 (10, 12)
fouchowensis, 194,196
fragilis, 6,121,122, 129,147,149,
153,290 (42, 43, 44)
fraudatrix group, 5, 44, 48, 29
fraudatrix, 73,274
fulleri, 58,287
fusca, 23
fuscanus, 11,12,13,15,18,20, 289
(2, 4)
fuscitarsis, 184,188
fuscocephala, 6,169,175,179, 182,
184,291 (66, 67, 68)
fuscosiphonis, 46, 48, 82, 87,113,
289 (26)
117, 129, 131
46, 48, 90,102, 110, 289
fuscus,
ganapathi,
(27)
gelidus subgroup, 6,171, 247
gelidus, 6,9,167,171,175, 181, 201,
247,291 (93, 94, 95)
geminus, 6,268, 288
gibbulus, 287
gnophodes, 241, 243
graminis, 129,131
habilitator, 165
hackeri, 286
halifaxii, 12,13, 17, 22,289 (3, 4, “ay
hayashii, 5,31, 286
hensemaeon, 194,196
Heptaphlebomyia, 163
Heteronycha, 163
hewitti, 287
hinglungensis, 5,34, 37,289 (12B)
hutchinsoni, 6,169,175, 181, 182,
188,291 (69, 70, 71)
impellens, 241, 243
incognitus, 288
incomptus, 46, 81,274, 290 (23)
inconspicua, 23
inculus, 287
inelegans, 184,186
infantulus, 5, 46, 49, 50, 58,290 (16)
infula, 258, 263
inornata, 117,129,131
jacksoni, 192,288
Jamesia, 11,12
Japanese encephalitis, 2,188, 245,
253
javanensis, 6,288
jenseni, 42,287
jepsoni, 241, 243,245
josephineae, 287
karatsuensis, 260,263
khazani, 5,275, 286
Laiomyia, 165
Lasioconops, 163
latifoliatus, 286
laureli, 42,286
lavatae, 287
Leucomyia, 163
litoralis, 236,239
longipes, 25,29
Lophoceraomyia, 2,3,5,6,8, 9,10,
11, 24, 33, 42, 43, 44, 119, 287,
289
Lophoceratomyia, 42
loricatus, 253,257
lucaris, 5,45, 79,290 (23A, B)
luridus, 13,15
luteola, 184,186
Lutzia, 3,5,8,9,10,11,12,13,289
macdonaldi, 47,50,61,69, 274, 290
(17)
macropus, 25
madagascariensis, 163
Maillotia, 23
malayi, 5,33, 34, 39, 42,289 (11, 12)
mammilifer group, 5, 44,50, 75,111
mammilifer subgroup, 5, 44, 47, 81,
111
mammilifer, 5, 45, 48, 83, 93,105,
290 (28)
microannulatus, 241,243
milni, 241,245
mimeticus, 192,219, 276,288
mimulus, 167,171,177,181, 202,213,
276,291 (80, 81, 82)
minimus, 184,186
minor, 5, 46, 48, 83, 95, 96, 275, 290
(29)
minutissimus, 271,287
Index to Culex of Thailand
Mochthogenes, 2,5,10,11, 24, 33,
42,43, 44, 286, 289
molestus, 200
mossmani, 215,217
multimaculosus, 17,22
navalis, 287
nematoides, 5,286
Neoculex, 2,3,5, 8,10, 11, 23, 33, 42,
43, 44, 286, 289
neolitoralis, 6, 8,167, 173,177, 236,
291 (89)
Neomelanoconion,
Nepenthes sp., 76
niger, 287
nigrescens, 39, 42
nigricephala, 241,243
nigropunctatus, 6,120,121,122,125,
133,145,291 (45, 46, 47)
nolledoi, 96, 98
obscurus, 286
ocellata, 260, 263
Oculeomyia, 165
okinawae, 5,286
omani, 236
orientalis, 288
Orthopodomyia sp., 117
pachecoi, 287
pallens, 200
pallidothorax, 6,120,121, 123,125,
127, 137,145, 163, 291 (48,
49, 50)
paludis, 241,243
papuensis, 6,121,122, 143,291 (51,
52, 53)
parainfantulus, 58,59
pathogens, 9
Pectinopalpus, 117
perplexus, 6,173,177, 182, 202, 205,
221,232,277, 291 (83)
peytoni, 45, 48,87, 92,99, 100,110, 290
30
117
Phalangomyia, 165
Philodendromyia, 42
pholeter, 45, 48, 103,290 (31)
pilifera, 23
pipiens group, 6,165,169, 182
pipiens, 163,198,275
pipiens pipiens, 275
pipiens quinquefasciatus, 6,9,12,17,
167,169, 173, 179, 182, 184,
192,291 (72, 73, 74)
2.95
plantaginis, 288
Plasmodium juxtanucleare, 169
Plasmodium vivax, 265
plegepennis, 253,257
pluvialis, 39
poicilipes, 163
propinquus, 288
Protomelanoconion, 33
Pseudoheptaphlebomyia, 163
pseudoinfula, 205,207 :
pseudosinensis, 6,171,177,179, 258,
265,274, 291 (101, 102)
pseudovishnui, 173,177, 182,205,
222, 232,277,291 (84)
pullus, 133,137
punctipes, 163
quadripalpis, 47, 49,50, 53, 65, 67,290
(18)
quinquefasciatus, 6,9,12,17, 167,
169, 173, 179, 182, 184, 192, 291
(72, 73, 74)
raptor, 17,18, 22
reesi, 194
reidi, 47,49, 66,290 (19)
vima, 117
roseni, 236
voubaudi, 65, 66
rubithoracis, 5,46, 49, 69,290 (20)
ryukyensis, 6,288
saibaii, 241, 243
salinus, 242
salsus, 241
salus, 241, 243
savawaki., 165, 260, 263
seanloni, 6,121,122,141, 147, 153,
291 (54, 55, 56)
sepositus, 268, 272
sericeus, 194,196
setulosus, 13
shebbearei, 6,288
shrivastavii, 288
siamensis, 227,230
simplex, 163
Simplicicornis, 5,286
sinensis, 6,171,175, 179, 258, 268,
292 (103, 104)
sitiens group, 6,169, 201
sitiens subgroup, 3,6, 236
sitiens, 6,167, 173,177, 181, 236,
239,292 (90, 91, 92)
solitarius, 288
somaliensis, 241,243
296 Index to Culex of Thailand
spathifurca, 6,120,121, 122, 151,
291 (57, 58, 59)
spiculosus, 46, 49, 87, 92,102, 105,
110, 155, 275, 290 (32)
spiculothorax, 6,120,122, 155,291
(60)
Spirogyra, 265
stigmatosoma, 165
stylifurcatus, 151,153
subalbatus (Armigeres), 117
sumatranus, 5,286
summorosus ; 225,230, 232
sylvestris, 65, 66
taeniarostris, 258, 263
taeniata, 55,58
taytayensis, 184,186
tenax (Leicester), 268, 272
tenax (Theobald), 260, 263
tenuipalpus, 5,24, 25, 29, 31, 289 (9)
termi, 6,120, 122,157, 290 (61, 62,
| 63)
territans, 23
Thaiomyia, 3,5,6,8,10,11,113,
119, 290
theileri, 275
Theobaldiomyia, 165
thurmanorum, 6,121,122,125, 127,
159,275,290 (64, 65)
tigripes, 12, 22
Transculicia, 165
traubi, 46, 49, 92,102,107, 108, 290
(33)
Trichopronomyia, 163
Trichorhynchomyia, 117
Trichorhynchus, 117
tricontus, 286
tripunctatus, 270,272
tritaeniorhynchus, 2,6, 9,167,169,
171,179, 181, 201, 209, 275,292
(85, 86, 87)
tuberis, 288
uniformis (Culex), 184,186
uniformis (Lophoceraomyia), 42,
275, 288
uniformis (Melanoconion), 25,29
univittatus, 276
Uranotaenia sp., 117
variatus, 47, 49, 50, 61, 69, 72, 274,
290 (21)
viridiventer, 6,163,275, 288
vishnui subgroup, 2,167,201, 202,
245,247
vishnui, 209, 225,276
vorax, 18,22
whartoni, 50,288
whitei, 173,177,182, 232, 233, 292
(88)
whitmorei, 6,171,175, 181, 247,
251,253,292 (96, 97)
whittingtoni, 236, 239
wilfredi, 45,48, 90,110,290 (34)
Wuchereria bancrofti, 9,120,155,
169, 201, 265
Wuchereria malayi, 143,245
Wuchereria sp., 209
yeageri, 42,286
Contributions
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Volume 2, Number 2, 1967
a, tad
THE SHORE BUGS (HEMIPTERA: SALDIDAE)
OF THE GREAT LAKES REGION
By Toby Schuh
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Volume 2, Number 2, 1967
ms
_ THE SHORE BUGS (HEMIPTERA: SALDIDAE)
OF THE GREAT LAKES REGION
By Toby Schuh
TABLE OF CONTENTS
Page
PTS ER eee Ne Mae Wee eRe wales eG EWU RO LG Red Ok ew Re adie 6 1
TNR PION ee icieek We Sik ae 0 ik lee ae bad a oes eek ah as 1
vols GBS EC 228 (ANESTRUS Jn) AGP ay Ue ARC ri FASTER PUNE aS Te YAO ra eg Pe er 1
De lS ee CaN Ae ee ey WE Guid Mew bee Wink ee ab bee ee ka oe 2
Te i is bee eee bee eee bee ee Me ete Gy icv Ligh) e Ne 3
PI EL se a eee Cae hie ay VW Ok eR EPE WO RHEE es 5)
PP Oe A Ce vis ip ete oC ee LR Gl IN Eas wee a hv oe celle HACER ei.
CPIM Peet wy 6 5 ho ce CE ERS OA GW MWR WR RW a Ma wiale oso bls e bac 9
Ee eee gy Wah Gi gen Wi Ge Soe Wk Gow Ow MVOC O EG Ue ROR ale eb wee els 10
GORUS WEEP Aaa oe se Ls ere i wiiesineains maces eles cne eee eee se 10
PE RC OL MORI ke cliW dio pad Oa Ae 6K SO eee wR ROR he eee 10
BCP CAN ie ATS Nokia Oh See bs Wad ee calce Hes ea Srce ak 11
PRC ORG Ae PE ag Ue eC elisa Sg ENS Da GOR Oe eR ENS Rees 11
SOEs ee eh se dae eR a Ges Gyles Boe OWE weds Wipe bee ws 12
ba UR Os Noe Ola aie kG eau dae be dienes vere 12
Te a a ce kb swe We no loo wy NN AMS, # eclaue wo OR cw Bie oe 14
ee hg cS ALG WH RE RO Le COL REO WO ee ee Sh ee ee 14
rete ONE Pas ee Wi oe BRE Wily ON G's We wie doe we OE Rein aa aes 16
ae ON Se ea aee Wa ace ee Ate oo OR oe ORD Ss we lee ees 16
NN EGE a he ike GA Mee ea ikta WwW Wo ee Daa Win awa eleibal a w acd is
ia eG aa 6 8 leva @ Wiel elk Wop rena awk Dome Rear eeees 18
ROU OF OUIAUN yk yikes che ROO Ak aise WED RURNU NS Wb be wen dig ee 19
AACE ILE IARI Ms NG yn Cua wae wh icele elem ka 6 20
i kg eis win ey be Siar ga kay WMA OLN 6 ce Wd ola eel ear a 20
ROPRR ta NS heh Co SRG NU ia (also ana apa GIN SRW nbn alla aisles ww bw wack 21
OO eee Pe aa ow ee ENS GA Soa NR Rall eG 21
CHOU TRL ln ls ek ys wie BV S/0 we week ee + 6.6 NE C4 Heh Raises Se wale eb oe 6 ae 22
Balcda: (Salda) anthracina ss ele ve eek le Oe Ss PADS GTEC De Mn rer ecnee ee 22
Galde (Saas DOUChervilten i. SEU Rie tae Deel wea Wea 0 23
Silca (ealan) WG! CU ue Cole Ce ee ee Ea es 24
Re Cy A iis bie a hE ne oe ee oe ee ee we wales a ) 24
Sala (Salda) GObSCura . os ies iw ee es ELE Ree Va greens 25
Salida { hamipracamtnia). CYABPICOMII B66 Sel be hea ek kee edie eles 26
BAlGa CT eloiedea) MelOONe cle heliaa sso weeks haw es ous Seameke en 26
LITERATURE CEEED oli vii w abiwle ea whinge oe bie alie ad eke a wo eee 27
THE SHORE BUGS (HEMIPTERA: SALDIDAE)
OF THE GREAT LAKES REGION
By
*k
Toby Schuh
ABSTRACT
Twenty species of shore bugs have been collected in the Great Lakes
region to date, and an additional six species might occur here. These species
represent five genera.
A discussion of the identification, habitat preference, and distribution is
given for each species. Distributional data is given in summary form for
well known species and complete data is given for species which are poorly
known or confused in the literature. Keys to all species occurring north of
the 36th parallel and east of the 100th meridian, with illustrations of 17
species, are included.
The Saldula "pallipes-palustris" group is discussed but no attempt has
been made to separate species. One new species, Saldoida turbaria is
described from Barry County, Michigan.
INTRODUCTION
This study is a review of the shore bugs of the Great Lakes region with
special emphasis on those species known to occur in Michigan. Material
used was collected during 1966 and supplemented with borrowed specimens.
Approximately 3000 specimens were examined, over 1500 of which were col-
lected expressly for this study. De
BIOLOGY
The shore bugs are semi-aquatic Hemiptera which commonly live along
the shores of ponds and streams; some live in marshy situations and some
are found well-removed from any permanent water. Saldids are character-
ized by long four-segmented antennae, a long three-segmented beak and long
legs. They are well adapted for living in littoral situations; the dense pubes-
cence on most species prevents wetting and holds a film of air around the in-
sect, allowing it to be submerged for short periods of time without harm.
*
Research conducted in the Dept. of Entomology, Michigan State University,
East Lansing, Michigan. Present address, Dept. of Zoology and Entomology,
University of Connecticut, Storrs, Connecticut.
2 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Shore bugs are predators and scavengers, feeding on dead or dying small
invertebrates. Ekblom (1926) observed them feeding on fly larvae burrowing
in the shore. Specimens I have observed in the laboratory were unable to
catch live insects but would readily feed on immobilized house flies. Saldula
ourayi Drake and Hottes, a common species around alkaline lakes in the
Great Basin, probably feeds on the larvae and adults of ephydrid flies which
are often very numerous along the shores of salty inland water bodies.
Oviposition in many species takes place in plant tissue and others lay
their eggs in the substrate. Wiley (1922) studied the life history of two North
American species and found they would readily oviposit in plant material in
the laboratory. The ovipositor is a heavily sclerotized sword-shaped organ,
with rows of teeth along its edge.
Many species are positively phototrophic, particularly the margin
dwellers. These often seek cover when the sun is obscured. The species
living in marshy areas and in dense vegetation are very seclusive and seldom
come out into bright sunlight. For many species there appears to be a
maximum and minimum substrate temperature at which activity begins and
ceases. The margin dwellers become active when the substrate reaches
about 600 F. and seek cover when the substrate temperature is above about
110° F.
Two different wing forms commonly occur in shore bugs. The fully
winged, or macropterous forms, have well-developed front and hind wings.
The short-winged, or brachypterous forms, have an abbreviated membrane
in the forewings and a reduction or loss of the hind wings. Most saldids are
active fliers, and nearly all are quick runners. The typical response when
alarmed is to jump and simultaneously begin flying. This escape method
makes most species difficult to collect. Distances traversed when alarmed
range from several inches in the flightless forms to several yards in the
flying ones.
TAXONOMY
Until recently the literature on the Saldidae was scattered, consisting
mostly of species descriptions. No comprehensive works have appeared
since Reuter (1912), except those of Cobben. Drake produced most of the
recent descriptions, but included no keys and very little ecological informa-
tion.
The taxonomic difficulties in the saldids are many, and several species
complexes exist. The range of morphological variation in some species is
extreme. The “eunomic series" for the hemelytra, as developed by Wagner
(1950), is helpful for separating species. It is based on the idea that the
pattern of pigmentation of the forewing is consistent within a species, varying
from the lightest to the darkest forms in a regular manner. |
An "eyespot" is often present medially along the lateral margin of the
endocorium (Figs. 1 & 13), particularly in the genera Saldula and Micracanthia.
Hemelytral pigmentation occurs in two basic forms: black pigment, encom-
passing most of the corium and clavus; and white pigment which forms marks
or spots. Areas lacking pigmentation usually appear translucent and are
clear to yellow, or light brown.
The systematic treatment used herein is that of Cobben (1959). Two
subfamilies are represented in the Great Lakes region, the Chiloxanthinae
Schuh: The Shore Bugs of the Great Lakes Region 3
and Saldinae. The Chiloxanthinae contains the genus Pentacora. The
Saldinae contains two tribes, the Chartoscirtini, including the genera
Micracanthia, Saldoida and Saldula, and the Saldini including the genus
Salda.
TECHNIQUES
The specialization of the shore bugs creates difficulties of collection and
identification. I have found the easiest way to collect them is with an aspira-
tor and a squeeze bottle of alcohol or water. The elusive nature of saldids
makes them difficult to pick up with the aspirator alone, but by first squirt-
ing them with alcohol or water they are wetted and knocked down and can be
sucked up before they are able to fly away. When there is a particularly
strong wind, sweeping with an aerial net is often a useful technique. By
simply walking along the shoreline and sweeping in front of you at the point
where the saldids begin to fly, large numbers of specimens can be obtained,
particularly if they are numerous. Coloration resembling the substrate
makes shore bugs difficult to see, and most collecting involves getting down
on the hands and knees and looking closely.
Best results are obtained if specimens are collected directly into 70-95%
ethyl alcohol. The dense vestiture of most species picks up dirt and debris,
and if collected dry they will often have to be cleaned before close examina-
tion can be made. All specimens should be mounted on card points with
"Elmer's" glue or a similar material.
When identifying specimens the characters of pubescence and wing mark-
ing must be observed with the light directed from the anterior end of the
specimen. Measurements in many cases require a micrometer eyepiece.
Measurements in the keys and text are of the greatest length or width of the
entire bug when viewed dorsally. Antennal ratios are indicated for some
species; these are noted as the relative lengths of the second, third and
fourth antennal segments, respectively (example: II-3, II[-2, IV-2). The
method of measuring the width of the vertex and one eye is indicated in
Figure 1.
Specimens collected for this study have hand-written numbers on the
printed locality labels. These numbers refer to my locality-habitat data
which will be on file in the Entomology Museum at Michigan State University.
I have retained some of these specimens, and deposited others in the collec-
tions of contributing individuals and museums; the majority of the specimens
are in the Michigan State University collection.
The data included in the "specimens examined" section of the text gives
abbreviations of the institutions from which the specimens were borrowed.
Specimens from my personal collection are denoted with a "TS".
ACKNOWLEDGEMENTS
I would like to thank the following institutions and curators for loans of
specimens (abbreviations used in the text are indicated in parentheses):
Field Museum of Natural History, Chicago, Illinois (FM), Henry Dybas;
Cornell University, Ithaca, New York (CU), L. L. Pechuman; Illinois Natu-
ral History Survey, Urbana, Illinois (INHS), Herbert H. Ross; Iowa State
4 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
University, Ames Iowa (ISU), Jean L. Laffoon; Michigan State University,
East Lansing, Michigan (MSU), Roland L. Fischer; Ohio State University,
Columbus, Ohio (OSU), Paul Freytag; University of Kansas, Snow Entomo-
logical Collections, Lawrence, Kansas (UK), George Byers; University of
Michigan, Ann Arbor, Michigan (UMMZ), Irving J. Cantrall; University of
Minnesota, St. Paul, Minnesota (UM), Ke Chung Kim; University of Wiscon-
sin, Madison, Wisconsin (UW), R. D. Shenefelt.
My special thanks go to: Mr. John T. Polhemus, Englewood, Colorado,
who identified some material, loaned and gave me numerous specimens, and
provided many ideas which have been incorporated in this paper (collection
abbreviation JTP); Dr. John D. Lattin, who initiated my interest in the
shore bugs; Dr. Frederick W. Stehr, who collected many specimens,
checked keys, and gave me much encouragement; Elwin D. Evans, who col-
lected many specimens which were very important in determining distribu-
tions; Richard J. Snider, whose help was invaluable in making the illustra-
tions; and Drs. T. Wayne Porter, Roland L. Fischer and Gordon E. Guyer
for their assistance in making this project a success.
Schuh: The Shore Bugs of the Great Lakes Region °.
KEY TO THE SPECIES OF SALDIDAE FROM THE GREAT LAKES REGION
NOTE: When examining dorsal pubescence the dorsum should be level
and the light should strike the specimen from the anterior.
5 (1)
6 (5)
7 (2)
Anterior lobe of pronotum with 2 prominent conical
tubercles (Figa. 5 GG) CiahOida) ois 4 sions Gales wees ee 5
Anterior lobe of pronotum without conical tubercles.............. 2
Membrane always with 5 visible closed cells (Fig. 2);
always macropterous( DeninCOra) 6s) 46d Mannie Wie soe eke os 7
Membrane with 4 visible closed cells if macropterous
(Fig. 1), or if brachypterous cells may not be visible........... 3
Hemelytra with long erect dorsal vestiture, at least as
long as the diameter of fourth antennal segment .............. 16
Hemelytra with short appressed pubescence, not longer
than smallest diameter of second antennal segment ............ 4
First cell of membrane produced anteriorly not more than
1/4 its total length beyond base of second cell (Figs.
YAR BG) dieunie ccna aracate ain tinanh ukeiaer MEA 6 ak apa nte eens ae aes 21
First cell of membrane produced anteriorly about 1/3
its total length beyond base of second cell (Figs.
Bh BO) iis a ew a Sis Wwe iis een ee i se ob 10
Pronotum deeply, concavely excavated on posterior margin
around scutellum; posterolateral corners of pronotum
NOt MPODUCEGS 5:59 iei6-2 4s a-aieid we bee Saldoida slossonae Osborn, p. 14
Posterior margin of pronotum not deeply, concavely
excavated, but nearly straight; posterolateral
corners.of pronotum prodived: (Fig, : 5) avin wmicnwedwas sees a oe 6
Posterolateral corners of pronotum sharply pointed;
hairs on eyes very short; vertex and frons with few
erect black hairs; hemelytra mostly ocherous and
QUEL as fs Pee be ea Saldoida cornuta Osborn, p. 14
Posterolateral corners of pronotum produced but not
sharply pointed (Figs. 5 & 6); hairs on eyes about as
long as diameter of base of second antennal segment;
vertex and frons covered with many long erect black
hairs; posterior half of hemelytra black, shining
(Wig By s cay wales iid ecae Saldoida turbaria new species, p. 12
Dorsum highly polished and shining, covered with erect
hairs at least twice as long as diameter of second
antennal segment; shaggy in appearance when viewed
from the side; dorsum almost devoid of shorter
appressed pubescence; pronotum, scutellum and
corium generally uniformly dark brown; lateral margin
of exocorium light; length over 5.0 mm...............0006
ih Gicduy Ce eee Go ale CHG eee Pentacora hirta (Say), p. 9
Dorsum not highly polished or shining, with erect
pubescence about the length of diameter of fourth
antennal segment, and/or short subappressed pubes-
cence; coloration variable, but not fitting above
JescripliONn vi bssace say ¥ ie ae GREG C504 BA WOES ores C48 ane ws 8
8 (7)
9 (8)
10 (4)
11(10)
12(10)
13(12)
14(13)
Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Lateral margin of pronotum and hemelytra with a row of
short spines; length of second antennal segment nearly
3 times that of third segment; large species, length
over OS mary Voce ey Pentacora signoreti (Guerin), p. 9
Pronotum without marginal row of short spines; second
antennal segment no more than 2.5 times as long as
CRIP SCOR WO ret ies od eee PCN cee ke we we ees 9
Dorsum clothed with erect hairs about as long as the
diameter of fourth antennal segment and shorter
reclining golden pubescence; antennae and legs with
short reclining pubescence; dorsum with partial light
transverse bands (Fig. 2); large species, length
ustially over 5/2 mm... ce ee oe Pentacora ligata (Say), p. 10
Dorsum with only short reclining pubescence, color of
dorsum uniformly light brown to mottled darker
brown; smaller species, length usually under
OOF ele erence. te Pentacora sphacelata (Uhler), p. 10
Dorsum black, sometimes with small scattered light
markings (Fig. 30); often brachypterous; body
especially convex when brachypterous ..........eeccescesves 12
Dorsum with mottled coloring (Figs. 3 & 4); head,
pronotum, scutellum and parts of corium dark,
exocorium and membrane mostly light, usually
macropterous; body TlAttened. oo Pee eee es See eee ees 11
Endocorium apically with a distinct unpigmented area
laterally (Pic. 3): length 4;0'to 6.0: mar... ee.
oer lea weeeeeres Salda (Teloleuca) pellucens (Fabricius), p. 26
Endocorium apically with only a small unpigmented
area (Fig. 4); leneth 870 te 90 Mere. ee eee ee
OTS eI Tt Salda (Teloleuca) bifasciata (Thompson), p. 22
Dorsum black, mostly shining, nearly devoid of
PUBESC CNC aoa 8 CS MELEE LE GERD GES GNSS EANA DOS See eee ees 13
Dorsum black, not shining, clothed with dense sub-
Bppressed PUDESCENCE Hs i. oo. EV eee oe eS ee eee ee eee cee 15
Second antennal segment black with a light stripe
dorsally; small slender species, length 3.8 to
0.6 mm, width 1.7 to 2.6 mm.; flattened in
appearance.......... Salda (Salda) obscura i walbaer). p. 25
Second antennal segment of nearly uniform coloration;
larger species with more robust and convex body
TOR SP er Ua he ee hee s as oe OE TG oe oe eee eee eas 14
Pronotum companulate (bell-shaped); antennae stout, |
segments 1, 3, and 4 dark, segment 2 lighter;
antennal proportions II-3, III-2, IV-2; length of
brachypterous forms 5.0°10'6.0' mm ..... 00.0030 0s... 0 os
ee ee ee ee Salda (Salda) anthracina Uhler, p. 22
Pronotum not particularly companulate; antennae slender,
all segments dark, proportions II-5, II-3, IV-3;
length of brachypterous forms 5.5 to 6.5 mm.,
macropterous forms 6.5 to 7.5 mm. (Fig. 30)...........
Se ie Pe oe: Salda (Salda) bouchervillei (Provancher), p. 23
15(12)
16 (3)
17(16)
18(17)
19(18)
20(17)
21 (4)
Schuh: The Shore Bugs of the Great Lakes Region 7
Dorsum with dense, reclining, golden pubescence; lateral
pronotal margins nearly straight, extending to pronotal
collar; sengee 4 0 10-51 OM Mint one bua Cade bein bee es
ee ee Salda (Salda) littoralis (Linnaeus), p. 22
Dorsum with very fine appressed pubescence, not generally
golden and shining; lateral pronotal margins terminating
abruptly before collar; length 5.5 to 6.5 mm..............
44 Wapeldepaie ted .. Salda (Salda) buenoi (McDunnough), p. 24
Hemelytra solid black, shining dorsally; antennal segments
one and 2, and legs light: antennal segments 3 and 4
black; anterior lobe of pronotum strongly swollen; length
3. O: £08 oD cit Wi: cis ou gemrulals a 4 greed tig Mh uate oO cub Sel adinieascneobia ds 6 «
evasliiaauabal oan onset Salda (Lampracanthia) crassicornis Uhler, p. 26
Hemelytra with variable markings, but not solid black;
antennae and legs variable in coloration; anterior lobe
of pronotum not..so- strongly swoOlleiic:. « o csissie sword steaierers 6 o's os 0 17
Eyes with numerous conspicuous hairs (visible at about
25X) nearly as long as diameter of second antennal
SOP TCI. sis eownneiw irra rede: a.anureueine deme Recaene Gatien ematisensie wie « 6 d-o'6 4. 18
Eyes Without. CONSPICUOUS AGT Sieii scare) x: sianeieeai aleine derargiorgcs ose 4s 6 6 20
Dorsum dull, nearly all black, with a marginal unpigmented
area near the apex of exocorium (Figs. 22 & 23); antennal
segments 2, 3 and 4 dark, covered with hairs nearly
twice as long as diameter of second antennal segment;
length. 3, 5:t0:4.2 MWe s6.6:d:is wees Saldula separata (Uhler), p. 21
Dorsum mostly dark colored; pronotum black, highly
polished; margin of exocorium mostly light (Figa:
VS 6o BO) cee cin bn aca an sain din ie ait ia a COO ei. ag 19
Body broad, ovate; pronotum broad, with a lateral
margins (Fig. 21); exocorium with 2 pruinose (bluish)
spots, endocorium with 3 pruinose spots, clavus with
one apical pruinose spot (Fig. 19); second antennal seg-
ment with hairs nearly as long as twice the diameter of
segments enetng.240 4.0 Wm. ok. ke a ko we ee be ke
2s Ls bee ee Wi eae. Saldula orbiculata (Uhler), p. 19
Body more slender, ovate; pronotum more narrowed ante-
riorly, with nearly straight lateral margins (Fig. 20);
clavus with one apical light spot; margin of exocorium
light (Fig. 18); second antennal segment with hairs about
as long as diameter of segment; length 3.2 to 4.0 mm.....
yp: ecceisitetiaten iss Ging see CSCC kw Ruteulaiiaa, abe Saldula severini Harris, p. 21
Hind tibia and second antennal segment with hairs sub-
equal in length to diameter of hind tibia; length 5.0
to 6.0 mam. Gvine pattern: Mig. 26) au a dicated, cise Senin acc d
«+ AAP WRUS dade als Bie tmat a ene Ae Saldula confluenta (Say), p. 16
Hind tibia and second antennal segment with hairs noticeably
longer than diameter of hind tibia; length 4.0 to 5.0 mm.
(wing pattern Fie. 27) +. 066 Saldula comatula (Parshley), p. 16
Length of second antennal segment equal to or greater than
distance across one eye and narrowest portion of vertex
(Fig. 1); large dark species with few markings on
hemetytra: LeNeiM-OVer 4.09 WAU 5 hoes Gey aciciwd Widlviabinis cm 6 22
22(21)
23(21)
24(23)
25(24)
26(24)
27(26)
Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Length of second antennal segment less than distance across
one eye and narrowest portion of vertex (Fig. 1); colora-
tion and markings variable; length seldom over 4.2 mm....... 23
Very large, robust, elongate-oval species, length 5.5 to
6.5 mm., width 2.5 to 3.2 mm.; dorsum covered with
short, reclining, shining, golden pubescence; mostly
dull black dorsally with scattered light markings;
anterior lobe of pronotum not particularly swollen,
transverse impression distinct but not deep............0.-
ie Oe ey VOR BE REE Salda (Salda) lugubris (Say), p. 24
More obovate and smaller than above, length 4.5 to 5.5
mm., width 2.0 to 2.5 mm.; dorsum with very neat,
short, appressed, dull silvery pubescence; dull black
dorsally with some light markings (Fig. 25); anterior
lobe of pronotum swollen (bulbous), distinctly sepa-
rated from head, and from posterior lobe by deep
transverse impression .......... Saldula nigrita Parshley, p. 17
Ovate; dark dorsally; eyespot usually fully developed;
pruinose (bluish) areas around eyespot and inner
margin of endocorium along clavus; without white "C"
shaped mark at apex of exocorium; lateral margin of
exocorium with light areas as in Fig. 13; length 3.0
tO. B.S Wie ea ere ARI Sw Saldula saltatoria (Linnaeus), p. 20
Generally obovate; no pruinose markings; length 2.5 to
4.5 mm.; "C" shaped mark present or absent .............6.4. 24
Hemelytra velvety black or brownish-black; endocorium
and clavus without distinct light markings (Figs. 11
G29) seme 2b 4000 2 MA. ies aes WHO Ses eee eee 25
Hemelytra not particularly velvety (color variable), or
if velvety then over 3.2 mm long; usually with dis-
tinct light markings on endocorium and clavus ...........eee8. 26
Lateral margin of endocorium with uninterrupted
light stripe of nearly uniform width (Fig. 11) .............
PAE NURS WON bee Micracanthia fennica Reuter, p. 10
Light lateral stripe not of uniform width and inter-
rupted posteriorly (Fig. 29)....Micracanthia ripula Drake, p. 11
Head, pronotum scutellum, clavus and endocorium mostly
black, partly shining, with short, scattered, silvery
pubescence; distinct white spot medially at lateral
margin of endocorium (Figs. 8-10 and 12); generally
small, compact forms, ovate; length 2.5 to 3.2 mm.......... 27
Dorsal facies variable; usually without distinct white
spot on endocorium as described above; larger
species, more obovate; length over 3.0 mm .......... 0.2020. 28
Exocorium mostly black with unpigmented areas inter-
rupting black subbasally and preeeicetly (Fig. 12);
Length 23 tO Se Mia a aoe Oe VV eR tres Ble wae ce oe
Ho ve be dee as Micracanthia quadrimaculata acRaNYpion). p. 10
Exocorium mostly unpigmented, with more or less well
developed black area medially (wing eunomy Figs.
8-10); length 2.5 to 3.5 mm... Micracanthia humilis (Say), p. 11
Schuh: The Shore Bugs of the Great Lakes Region
28(26) Dorsum mostly black; exocorium interrupted by two unpig-
mented areas laterally; a more or less well-developed
"C" shaped white mark apically at mesal margin of
exocorium (Fig. 28); length 3.0 to 4.0 mm...............
aks ee Se CS Micracanthia floridana Drake and Chapman, p. 10
Dorsum variable; exocorium with unpigmented areas not
as-above,. and lacking "CC" iaped Marko. cs ive eee ees ce’s 29
29(28) Lateral margin of exocorium never interrupted with dark
markings; corium with three distinct whitish spots
(see discussion, Fig. 24); dorsum with short appressed
golden pubescence; length 3.2 to 4.0 mm.................
cS Mower er a Gee Mere Saldula opacula (Zetterstedt), p. 18
Lateral margin of exocorium usually interrupted with
dark markings; white pigmented areas not distinct
as above; ledeth 3.05192, 9 tits ool ee a Bei. bane. ad 30
30(29) Pronotum sometimes with a light lateral margin; corium,
membrane and apex of clavus usually light; lateral
margin of exocorium with more or less well-developed
dark areas apically and posterior to middle; a broad
transverse "“orange-colored" darkened area at apex of
corium when viewed from a distance (wing eunomy Figs.
L4<L7)i lengthy 3.240455 mani oes Fs Oe a
SOURE LENG ee te eee Saldula ablusa Drake and Hottes, p. 14
Pronotum never with light lateral margin; hemelytra from
almost completely light to very dark; exocorium with
darkened area apically (indistinct in very light speci-
mens), and 2 dark areas developing parallel to lateral
margin but not touching margin except in darker
specimens; development of markings in exocorium
accompanied by darkening of endocorium; without
“orange-colored" transverse band at apex of corium
(see discussion); length 3.0 to 4.5 mm............ccceees
bse Sst algta ee eee »....... oaldula "pallipes-palustris" group, p. 20
SUBFAMILY CHILOXANTHINAE
Pentacora Reuter
The species of the genus Pentacora are primarily halophilous. Four
species occur in North America, of which one, Pentacora ligata (Say), has
been collected in the Great Lakes region. The other three species are
apparently largely restricted to the coastal areas, but some inland records
are known, particularly from areas in the Mississippi River drainage (see
Drake, 1949).
Pentacora hirta (Say) has been recorded from most of the Atlantic
and Gulf Coast states as well as Indiana and Iowa (Drake, 1949). Pentacora
signoreti (Guerin) shows a similar distribution, but is also known from the
southern California coast and the Salton Sea. This species is usually found
in saline situations, and is recorded by Froeschner (1962) from mineral
springs in Boone and Jefferson Counties, Missouri. I have examined
specimens of signoreti in the University of Michigan Museum of Zoology
10 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
from Devil's Lake, North Dakota.
Pentacora sphacelata (Uhler) is the most restricted in habitat, occurring
only along the coast, but it has the widest range. It occurs along the coast
from Massachusetts to the Gulf Coast, southern California, the Caribbean,
Peru, and the Mediterranean (Cobben, 1960b).
Pentacora ligata (Say), 1831
Figure 2
Discussion--Pentacora ligata has a dense, semi-erect dorsal vestiture
which is not nearly as long as that in hirta; the dorsal vestiture of spacelata
is shorter than in ligata and more appressed. The dorsum of ligata is only
slightly shining as compared to the highly polished dorsum of hirta. The
second antennal segment is no more than 2.5 times as long as the third in
ligata, which will separate it from signoreti in which the second segment is
nearly three times as long as the third. |
Habitat preference--Pentacora ligata is a large and very agile species
generally found on logs or large rocks in streams, and occasionally around
standing water. It is often found in association with Saldula confluenta (Say).
Distribution--Records are known for most of the states east of the
Mississippi River and for eastern Canada. I examined 59 specimens from
Michigan collected between June 18 and September 17 and 111 specimens from
Illinois, Indiana, Minnesota, New York, Ohio, Tennessee, Wisconsin, and
Ontario collected between June 5 and September 1.
SUBFAMILY SALDINAE
TRIBE CHARTOSCIRTINI
Micracanthia Reuter
This genus is holarctic and includes species occurring both in Europe and
North America. Micracanthia fennica Reuter (Fig. 11) occurs in Europe
and is known from Oregon, Colorado and Nebraska; Micracanthia quadri-
maculata (Champion) (Fig. 12) occurs in the western United States, extending
into Central America, and I have seen specimens from as far east as South
Dakota. These species have been illustrated and included in the keys.
Several species of Micracanthia occur in the Gulf Coast region. Among
these are pumpila Blatchley, husseyi Drake and Chapman, and drakei Cobben
which seem to be closely allied to humilis (Say) (see Cobben, 1960b, and
Drake and Chapman, 1952, for discussion of these species).
Micracanthia floridana Drake and Chapman, 1953
Figure 28
Discussion--One of the largest species in the genus, Micracanthia flori-
dana looks much like some of the Saldula species. The "C" shaped mark at
the apex of the exocorium separates floridana from other members of the
genus.
Schuh: The Shore Bugs of the Great Lakes Region 11
Habitat preference--Micracanthia floridana is usually found on logs or
rocks and other objects hanging over streams or standing water. In the
Great Lakes region associated species include Saldula confluenta and pos-
sibly Pentacora ligata.
Distribution--Originally described from specimens collected in Florida,
Mississippi and Colorado (Drake and Chapman, 1953) records from Michigan
represent an extension of the known range of this species.
Specimens examined--MICHIGAN: Emmet Co., Maple R. at Brutus Rd.,
T36N R4W Sec 24-25, 23 Jul 1966, T. Schuh, E. Evans, 10°; Wexford Co.,
Sherman, Manistee R., T24N R24W Sec 31, 5 Sept 1966, E. Evans, 4c, 29.
FLORIDA: St. Augustine, Jan 1901, W. J. Gerhard, 19 (FM). ILLINOIS:
Havana, Quiver L., Station F., 28 July 1894, 19 (INHS). NEW JERSEY:
Camden, 1X: 18, S. T. Kemp, 19 (FM).
Micracanthia humilis (Say), 1831
Figures 8-10
Discussion--Micracanthia humilis is quite variable in size, ranging from
2.9 to 3.5 mm. in length. It is however rather uniform with respect to the
pigmentation of the hemelytra (wing eunomy, Figs. 8-10). The white spot at
the middle of the lateral margin of the endocorium and the elongate black area
in the exocorium will separate humilis from other members of the genus
occurring in the Great Lakes region. Cobben (1960b) and Drake and Chapman
(1952) discuss closely allied species occurring in the Gulf Coast region.
Habitat preference--Damp sandy stream and pond margins are the most
common habitats of humilis. It is usually found somewhat removed from the
water in areas with small amounts of grasses and sedges. Species collected
in association with humilis were Saldula ablusa Drake and Hottes and Saldula
"pallipes-palustris".
Distribution--Micracanthia humilis is the most common member of the
genus in eastern North America. It ranges from southern Canada into the
Caribbean, and from the Atlantic coast west to the Rocky Mountains. Ihave
examined 365 specimens from Michigan, Georgia, Illinois, Indiana, Iowa,
Minnesota, North Carolina, North Dakota, Ohio, Tennessee, Wisconsin,
and Florida; collection dates range from April 6 to October 21.
Micracanthia ripula Drake, 1958
Figure 29
Discussion--Micracanthia ripula is rather dark with heavy black pigmen-
tation on the corium. The dorsum in general appears velvety brownish-black
because of the thick pubescence. As with most members of the genus, a
light area is generally present at the posterior end of the eyespot. The mar-
gin of the exocorium is light, interrupted submedially with black. A distinct
_ light spot is present near the apex of the exocorium. Micracanthia humilis
in the darker forms (Fig. 10) closely resembles ripula, but usually has a
more well-developed eyespot and silvery, less dense dorsal vestiture; the
dorsum of humilis is black and almost shining. The membrane of humilis
is uniformly light, whereas the membrane of ripula is usually darker and
blends into the corium anteriorly.
12 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Habitat preference--Seven specimens were collected from two localities
in Keweenaw County, Michigan. Both areas had a sandy substrate and some
vegetative cover. This species seems to seek cover and darts out period-
ically. At a location near Lake Superior I found ripula in small clumps of
moss in association with Salda bouchervillei (Provancher) and Salda obscura
(Provancher). As is typical of M. quadrimaculata and humilis, ripula seeks
damp to almost dry sandy substrates.
Distribution--Described from Churchill, Manitoba (Drake, 1958), ripula
has also been recorded from Marble Canyon, Lilooet, and Kamloops, British
Columbia (Scudder, 1961).
Specimens examined--MICHIGAN: Keeweenaw Co., Montreal R., T58N
R30W Sec 14, 10 Jul 1966, T. Schuh, F. W. Stehr, 10" eek Keweenaw Co.,
TOTN R380W Sec 36, 10 Sul 1966, T. Schuh, F. W. ‘Stehr, A |
Saldoida Osborn
Osborn (1901) erected the genus Saldoida to accept two North American
species, cornuta Osborn and slossonae Osborn. Reuter (1912) placed this
* genus in a separate subfamily, the Saldoidinae. Drake and Chapman (1958)
review Saldoida. Cobben (1959) ee? Saldoida in the Saldinae, under the
tribe Chartoscirtini.
Saldoida turbaria, new species
Figures 5-7
Description--Small, slender, broadened posteriorly; mostly black with
brown and white markings on the hemelytra; brachypterous.
Head--Black, shining, collar-like behind eyes; eyes conspicuously
bulging; ocelli yellow, located at middle of vertex, separated by approxi-
mately the width of an ocellus; rostrum, labrum and-tylus testaceous;
rostrum extending between hind coxae; vertex and frons with numerous long,
erect, black hairs; eyes conspicuously hairy under moderate magnification
(25X).
Thorax--Black, shining, anteriorly with a distinct collar; lateral margins
nearly straight with posterolateral angles produced, angulate; posterior mar-
gin straight at corners, shallowly and evenly concave around base of scutellum;
dorsal surface of anterior lobe of pronotum elevated to form a pair of large
erect conical tubercles, nearly as long as width of one eye measured across
the vertex; dorsal surface of pronotum covered with erect black hairs;
scutellum black, shining, with decumbent golden pubescence; posterior por-
tion of scutellum elevated, glabrous.
Abdomen--Venter black, shining; second sternite densely clothed with
decumbent golden pubescence; sternites three through six with scattered
semi-erect golden pubescence; see Figure 6 for form of sixth abdominal
sternite; ovipositor barely visible at tip of abdomen in ventral view.
Wings--Hemelytra reduced, brachypterous; sparsely covered with long,
erect, black hairs, at least as long as the greatest diameter of the third
antennal segment; outer margin of clavus dull black, inner margin ferrugi-
neous; corium mostly fuscous with some lighter, nearly white areas; mem-
brane and posterior part of corium shining black; membrane without visible
venation; embolium whitish; margin around membrane nearly transparent.
Schuh: The Shore Bugs of the Great Lakes Region 13
Extremities--Antennae long, about two-thirds the length of body; seg-
ments one and two slender, flavous; segment two white apically, one and two
with short, dark, decumbent, pubescence; segment three greatly enlarged,
at least twice the diameter of segment two; segment four slightly smaller in
diameter than three; segment 3 entirely black, segment 4 black basally and
white apically; segments three and four with decumbent golden pubescence
and longer, less dense, semi-erect black hairs.
Legs testaceous; coxae and basal half of femora light, apical portion of
femora and entire tibia darker; all legs with short decumbent golden pubes-
cence, all tarsi with short dark spines.
Measurements--Body length 3.2 mm.; body width 1.5mm. Antennal
proportions: I-35; I-60; I-60; IV-60.
Holotype--Brachypterous female; type locality: Michigan, Barry County,
Purdy Bog, 3 miles ESE of Hickory Corners, TIN, R9W, Section 36, 29 July
1966, Toby Schuh Collector. Deposited in the Michigan State University
Entomology Museum.
Paratype--Brachypterous female; same locality and collection data as
holotype. In the collection of the author.
Discussion--This species, though described from only two female speci-
mens, was deemed distinct enough to be given specific status. Comparison
with specimens of Saldoida cornuta Osborn, which appears to be the most
closely related species, indicates several important taxanomic differences.
The eyes of turbaria are visibly hairy under moderate magnification (25X),
the hairs being nearly as long as the diameter of the base of the second
antennal segment; Saldoida cornuta has much shorter hairs on the eyes. The
vestiture of the dorsum, especially the hemelytra, is distinct, being long,
black, and erect, whereas in cornuta it is much shorter and more appressed.
Other characters such as the coloration of the hemelytra and the antennae
should eliminate any confusion with previously described species.
This new species is designated as "turbaria" to indicate its occurrence
in a bog.
Habitat preference--Previous collections of Saldoida in the southeastern
United States have been from low marshy areas where sundew plants and
ecologically similar types of vegetation grow. The bog in the Great Lakes
region is the situation which most closely approximates this type of habitat.
Correspondence with Dr. John D. Lattin indicates that a nymph of the genus
Saldoida was collected at Mud Lake, a bog, in Cheboygan County Michigan,
in 1951. This specimen was secured from a burlese funnel sample while
processing Sphagnum for Ceratocombus (Dipsocoridae). The nymph could
not be identified to species and no adults were found by Lattin. I made col-
lections at Mud Lake in July, 1966, and many large moss samples were
taken, but no Saldoida were obtained. Later in July, the specimens herein
described were obtained from Purdy Bog, in Barry County, Michigan. Be-
cause of its small size, secretive habits, and the difficulty of collecting in
the bog mat, no additional specimens were obtained.
Saldoida turbaria was collected near the edge of the bog mat on a small
muddy margin of the bog lake. The area was protected by higher vegetation
_ growing at the mat edge, and received very little sunlight.
Numerous specimens of Antillocoris minutus (Bergroth) (Lygaeidae) and
and Hebrus burmeisteri Lethierry and Severin (Hebridae) were collected in
the Sphagnum in the area where turbaria was taken.
14 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Behavior--Saldoida turbaria is not agile. Movements are jerky, giving
the appearance of walking one leg at a time when the bug is crawling on the
uneven mossy substrate. Short jumps of about one inch are made when the
bug is disturbed. The antennae are in constant motion, being moved up and
down alternately; this movement of the large antennae helps to locate the
insects since they are very small and prefer shaded areas.
The legs are used by the insect to clean itself. The front legs are peri-
odically brushed over the antennae. The two pairs of hind legs are rubbed
against each other, especially the tibiae, and the hind legs are raised over
the abdomen and stroked posteriorly. The bug periodically interrupts its
wanderings and performs these cleaning motions.
Distribution--The genus Saldoida was previously considered to be pri-
marily tropical and subtropical. The two species recorded from the United
States, cornuta and slossonae, are confined mainly to the Gulf Coast region
and the southern Atlantic Coast of the United States. The most northern col-
lection records are for slossonae from Tuckahoe and Helmetta, New Jersey
(Drake and Chapman, 1958). Saldoida armata Horvath is known from Taiwan
and the Phillipines. Saldoida turbaria represents an extension of the known
range of the genus in North America. The distribution of this new species
should be roughly coincident with that of bogs in the Great Lakes region.
Saldula Van Duzee
Saldula is a cosmopolitan genus, and the largest and most diverse in the
Saldidae. Eleven species are known from northeastern North America. The
"pallipes-palustris" group contains examples which closely resemble the
European species, Saldula pallipes (Fabricius) and palustris (Douglas).
Separation of the North American forms of this group is extremely difficult
and there is uncertainty as to the actual names which should apply to the
New World forms.
Saldula ablusa Drake and Hottes, 1954
Figures 14-17
Discussion--Saldula ablusa appears to be very closely related to the
Saldula "pallipes-palustris" group, and the very light forms of the two are
often difficult to separate. Saldula ablusa has a somewhat broader pronotum
and dorsally is covered with more golden pubescence than "pallipes-
palustris". The pronotum often has a light lateral margin, a character not
occurring in "pallipes-palustris". The light margin may be completely
lacking or may be developed to the point where the pronotum is almost com-
pletely light posteriorly. Most specimens have light hemelytra, the darkest
forms being about as dark as a moderately dark "pallipes-palustris". The
hemelytra have an "orange" coloration and are usually opaque, whereas
"pallipes-palustris" is usually yellowish in the light forms and the hemelytra
are more translucent. When viewed from a distance, a darkened transverse
band is apparent on the posterior part of the corium; this does not appear in
"pallipes-palustris". The eyespot in ablusa is seldom outlined with dark
pigment, but is set off by what appears to be a whitening of the outside ring
against the general background coloration of the wing; "pallipes-palustris"
begins to show a dark outline of the eyespot in the forms which are not
Schuh: The Shore Bugs of the Great Lakes Region _ 15
heavily pigmented. A dark spot at the lateral margin of the hemelytra about
two thirds of the way posterior on the exocorium is almost always well-
developed, and occasionally a pigmented line appears anterior to the spot in
the middle of the exocorium; "pallipes-palustris" has the lateral spot with-
drawn from the wing margin and it does not become enlarged except in the
darker forms. The second antennal segment varies from light apically to
completely light; this character is usually more constant in "pallipes-
palustris", the second segment being light apically and darkened elsewhere,
with the entire segment only rarely entirely light. Refer to figures 14
through 17. |
Habitat preference--Saldula ablusa occupies a habitat similar to that of
"pallipes-palustris" and is most commonly found in association with it. All
_ the collections I have made in Michigan were from the sandy shores of ponds
and streams. The substrate is usually light colored, smooth, damp sand;
most localities have a limited amount of scattered vegetation. The numbers
of ablusa in a given habitat do not seem to be as great as those of "pallipes-
palustris". The largest collection of ablusa was ten specimens from the
Otter River in Baraga County, Michigan; about an equal number of "pallipes-
palustris" were taken from this locality. Most collections show about a ten
to one ratio of "pallipes-palustris" to ablusa. As with other species, ablusa
is usually colored very similarly to the substrate, and they seem to seek an
area in which they are best concealed. Micracanthia humilis was collected
in association with ablusa.
Distribution--Saldula ablusa was described from Sand Point, Huron
County, Michigan (Drake and Hottes, 1954). The distribution seems to be
restricted primarily to the northeastern United States and Canada, but I
have seen specimens from the northwestern United States which look very
much like ablusa. Other similar species, such as Saldula opiparia Drake
and Hottes occur in the western United States and more work will have to be
done to determine if ablusa actually occurs in the west.
Specimens examined--MICHIGAN: Alger Co., Kingston L., W. of
Grand Morais, 15 Jun 1966, F. W. Stehr, 50’, 72; Baraga Co., Otter R.,
TOIN R384W Sec 9, 10 Jul 1966, T. Schuh, F. W. Stehr, 40°, 69; Benzie Co.,
Honor, 29 May 1966, E. Evans, 50°, 69; Cedarville, 14 Jul 1955, G. E.
Guyer, 20° (MSU); Cheboygan Co., (30°, 62 from Douglas Lake area from
3 Jul to 30 Jul), (UK); Cheboygan Co., Douglas L., 4 Jul 1923, H. B.
Hungerford, 12 (JTP); Cheboygan Co., Douglas L., 29 Jun 1923, H. B.
Hungerford, 19 (JTP); Clinton Co., Park L., 10 Apr 1966, T. Hlavac,
19; Dickinson Co., Sturgeon R., T40N R28W Sec 23, 8 Jul 1966, T. Schuh,
1c’, 19; Emmet Co., Crooked L. at Conway, T35N R5W Sec 13, 23 Jul 1966,
T. Schuh, E. Evans, 1c; Emmet Co., 14 Jul 1935, M. Sanderson, 1c’, 19
(JTP); Grand Traverse Co., Boardman R., T26N R11W Sec 19, 22 Jul 1966,
T. Schuh, E. Evans, 10°; Hog Island, L. Michigan, 9 Jul 1939, H. B.
Hungerford, 10 (UK); Houghton Co., Crystal L., T47N R36W Sec 2, 10 Jul
1966, T. Schuh, 12; Ingham Co., 24 Oct 1949, H. C. Chapman, 1c", 19
(MSU); Keweenaw Co., Eagle R. at L. Superior, T58N R31W Sec 19, 9 Jul
1966, T. Schuh, F. W. Stehr, 19; Mackinac Co., Cut R. at L. Michigan,
17 Jun 1966, F. W. Stehr, 1c‘, 12; Menominee Co., L. Michigan, T34N
R25W Sec 20, 17 Jun 1966, F. W. Stehr, 1c’, 49; Ottawa Co., L. Macatawa,
TON R1I6W Sec 27, 24 Jun 1966, T. Schuh, 42; Roscommon Co., 24 Jun 1951,
R. R. Dreisbach, 20°, 12 (MSU); Schoolcraft Co., Clark's Ditch, 2 mi. W
Seney, 8 Jul 1966, F. W. Stehr, 1c‘, 19. ILLINOIS: Waukegan (beach),
16 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
23 Aug 1906, 4c’, 119 (INHS). INDIANA: Clark Jct., 21 May 1905, W. J.
Gerhard, 1c, 39 (FM); Clark Jct., 28 May 1905, W. J. Gerhard, 39 (FM).
MINNESOTA: 3 mi. S. Afton, 5 May 1951, D. Scharff, 60°, 29 (UM); Ball
Club, Ball Club R., 3 Jul 1935, D. G. Denning, 10° (UM); Itasca St. Pk.,
26 Jun 1958, L. R. Abra., 10° (UM); Pine Co., Mouth Snake R., 16 May
1959, 10° (UM); Ottawa, 18-19 Jul 1922, W. E. Hoffman, 19 (UM); Swan R.,
2 Jul 1935, D. G. Denning, 12 (UM); Washington Co., Afton, 9 May 1959,
22 (UM). NEW YORK: Nassau Co., Jones Beach St. Pk., 28 Jun 1959,
G. C. Eickwort, 12 (MSU). OHIO: Cleveland, 29 Jun 1934, M. W.
Sanderson, 19 (UK). SOUTH DAKOTA: Custer St. Pk., 20 Jun 1955,
R. L. Fischer, 1o’ (MSU). CANADA: Ontario, Prince Edward Co.,
Brimely, 24 Jul 1932, 192 (JTP).
Saldula comatula (Parshley), 1921
Figure 27
Discussion--Although this is primarily a western species I have included
it to make the keys more complete. Saldula comatula and S. confluenta (Say)
are the only "hairy" species which lack conspicuous hairs on the compound
eyes. The wing pigmentation pattern and long hair on the second antennal
segment and tibiae make comatula an easily recognized species. One western
species, S. pexa Drake, resembles comatula very closely but has a slightly
different wing pigmentation pattern and lacks the shaggy antennae and legs.
Usinger (1956, p. 224) illustrates a specimen labeled as comatula, which is
actually pexa.
Habitat preference--The habitat of comatula is unlike any of the other
eastern "hairy" species in that it is generally found on open stream and pond
margins of a muddy or sandy nature where little vegetation is present. I
have collected this species in the west and have never found it in large numbers,
but have found that it occurs with regularity along most pond and stream mar-
gins where the "pallipes-palustris" group is present.
Distribution--Originally described from British Columbia (Parshley,
1921), the known range of comatula now covers most all of the United States
and Canada west of the Mississippi River. Harris (1937) records comatula
from Batesland, South Dakota. This species might be found in the Great
Lakes region when more intensive collecting is done.
Specimens examined--COLORADO: Dolores, 15 Aug 1925, C. J. Drake,
19 (UW). ILLINOIS: Illinois, Andreas Bolter Collection, 19 (INHS).
NEBRASKA: Morrill Co., 22 Aug 1951, R. R. Dreisbach, 1c‘ (MSU).
SOUTH DAKOTA: Custer St. Pk., 20 Jun 1955, W. A. Drew, 19 (MSU).
Saldula confluenta (Say), 1831
Figure 26
Discussion--Saldula confluenta can be easily distinguished from all other
members of the genus by its size, as it is usually over 5 mm. long, whereas
most other species seldom exceed 4.5 mm. The long, erect dorsal vestiture
and rather distinctive coloration pattern of the hemelytra are diagnostic.
Habitat preference--Saldula confluenta is usually found on rocks and logs
in streams, or on rocks on the stream margin. Many fifth instar nymphs
Schuh: The Shore Bugs of the Great Lakes Region 17
were running on a partially shaded black muddy bank of the Looking Glass
River in Clinton County, Michigan in early June. No adults were collected
at this locality but nymphs were reared through to adults in the laboratory
in about four days. Saldula confluenta is seldom very numerous which makes
collecting large numbers of specimens difficult. Little mention has been
made of this species in the literature, probably because of its rather special-
ized habitat and great agility, both of which make it difficult to collect, and
consequently rather uncommon in collections. Pentacora ligata is often col-
lected in association with confluenta.
Distribution--Collection records indicate confluenta occurs over most of
the northeastern United States, and I have taken specimens from all parts of
Michigan. The most southerly and westerly record is from Texas (Froeschner,
1962). I have examined 63 specimens from Michigan collected between
June 12 and September 8, and 25 specimens from Illinois, Minnesota, New
York, Ohio, North Carolina, Tennessee, Wisconsin and Ontario collected
between May 29 and August 8.
Saldula nigrita Parshley, 1921
Figure 25
Discussion--Typically quite large, Saldula nigrita ranges in length from
4.3 to 5.5mm. Two forms seem to exist with respect to wing pigmentation.
The most common type is dark, the corium being marked with a number of
light spots; this is the only form that I have collected in Michigan. I have
collected a lighter form in the Great Basin, the hemelytra of which are un-
pigmented except for the veins. The legs of Saldula nigrita are almost en-
tirely black except for a light dorsal stripe on the femur. The second
antennal segment is relatively longer than in closely related species (anten-
nal proportions: I-19; II-12; IV-11). The relative length of the antennal
segments and the dark legs will separate nigrita from the "pallipes-
palustris" group. The dorsum is neat in appearance with only a small
amount of silvery pubescence. Saldula nigrita might be confused with Salda
lugubris (Say) but is much more slender and more obovate.
Habitat preference--Saldula nigrita has a rather specialized and well-
defined habitat. Chapman (1962) notes nigrita occurring on stones in streams
and on the shore; Lindberg (1958) collected it on sandy and stony banks of
streams, and on pond and lake shores. At the two locations where this
species was taken in Michigan, I found it on exposed rocks in streams and
along stream margins. Saldula nigrita has a habit of crawling under rocks
and then appearing momentarily, but never seems to sit in one exposed
place on the substrate for an extended period of time.
Distribution--Saldula nigrita was described from Duncan, British
Columbia (Parshley, 1921). Subsequent collecting has greatly extended the
known range, indicating a boreal distribution. Records are known from
Newfoundland (Lindberg, 1958) and as far south in the Rocky Mountain
system as Carlsbad, New Mexico. The distribution pattern of nigrita is
fairly common for the saldids of North America, particularly the genus
Salda. Records for nigrita are known from stream mouths along the Pacific
Coast in Oregon, and although this does not appear to fit the general distri-
bution, the coastal climate is cool most of the year and not unlike the climatic
conditions encountered in the mountains and northern latitudes in the summer.
18 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Specimens examined--MICHIGAN: Baraga Co., Otter R., T51N R34W
Sec 9, 10 Jul 1966, T. Schuh, F. W. Stehr, 310°, 209; Dickinson Co.,
Sturgeon R., T29N R28W Sec 13, 12 Jul 1966, T. Schuh, 40’, 69.
MINNESOTA: Grand Marais, Devils Track R., 7 Aug 1927, W. C. Stehr,
60°, 62 (UM).
Saldula opacula (Zetterstedt), 1840
Figure 24
Discussion--Saldula opacula closely resembles the "pallipes-palustris"
complex, but can be separated rather easily when specimens are available
for comparison. No dark pigmentation is present in the lateral portion of
the exocorium. Some forms of “pallipes-palustris" can be confused with
opacula because of the light wing margin, but the development of this light
area is of a different nature than in opacula. The eyespot is always evident,
the posterior portion being clouded with white, the anterior portion yellowish.
The posterior part of the eyespot of "pallipes-palustris" does not have white
pigmentation. A distinct white area occurs near the apex of the endocorium,
just above the base of the second cell. An elongate, oval, white area is
always evident subapically in the exocorium, lying mesad of the dark pig-
mented line which parallels the wing margin. In the typical pigmentation
pattern the veins in the endocorium darken rather uniformly as the stripe
paralleling the wing margin develops.
Habitat preference--A holarctic species originally described from
Europe, opacula inhabits very wet places such as marshes and bogs.
Lindberg (1958) notes collecting opacula from Sphagnum; I collected four
specimens from the mossy banks of a stream in Kalkaska County Michigan.
This species is seldom found in large numbers.
Distribution--The distribution of opacula is similar to that of Saldula
saltatoria (Linnaeus) and apparently occurs over most of Canada and the
northern United States. 3
Specimens examined--MICHIGAN: Baraga Co., Keweenaw Bay, T50N
R33W Sec 1, 10 Jul 1966, T. Schuh, F. W. Stehr, 19; Cheboygan Co.,
29 Jul 1918, R. F. Hussey, 10° (UMMZ); Cheboygan Co., 23 Jul 1955, R. R.
Dreisbach, 19 (MSU); Cheboygan Co., 23 Jul 1952, H. B. Hungerford, 1c,
1? (UK); Emmet Co., Petoskey, T35N R5W Sec 34, 23 Jul 1966, T. Schuh,
E. Evans, 12 Huron Co., Point aux Barques, 17 June 1922, R. F. Hussey,
1? (UMMZ); Ingham Co., East Lansing, 26 Sept 1965, E. Evans, 10° (MSU)
Ingham Co., East Lansing, 2 Nov 1963, R. B. Willson, 19 (MSU);
Kalamazoo Co., Gull Lake Bio. Sta., 25 Jul to 7 Aug, 59, 20° (MSU);
Kalkaska Co., Desmond Crk., T28N R8W Sec 28, 22 Jul 1966, T. Schuh,
E. Evans, 20°, 29; Livingston Co., E. S. George Reserve, 20 March 1938,
J. J. Friauf, 22 (UMMZ); Mackinac Co., Cut R. at L. Michigan, 17 Jun
1966, F. W. Stehr, 10°; Ontonogan Co., Sooty Crk., T50N R44W Sec 24,
11 Jul 1966, T. Schuh, 10‘; Van Buren Co., 8 May 1949, H. C. Chapman,
30°, 192 (MSU). INDIANA: Clarke Jct., 7 Aug 1904, W. J. Gerhard, 22
(FM). MINNESOTA: Crookston (light trap), 19 Jul 1935, D. G. Denning,
129 (UM); Glenwood Park, 14 Aug 1925, G. O. Wiley, 19 (UM); Lancaster,
25 Jun 1937, D. G. Denning, 19 (UM); Mille Lacs Co., Mille Lacs, 18 May
1940, C. E. Mickel, 20, 12 (UM); Ottawa, 18-19 Jul 1922, W. E. Hoffman,
1? (UM); Ramsey Co., Golf Ponds, 1 Jun 1923, W. E. Hoffman, 1c", 19
Schuh: The Shore Bugs of the Great Lakes Region 19
(UM); St. Anthony Park, 9 Jul 1923, H. H. Knight, 22 (UM); St. Paul, 1 Jul
1923, G. O. Wiley, 12 (UM); NEW YORK: Buffalo, Sept 1886, E. P. Van
Duzee, 19 (ISU); Ithaca (at light), 1 Jul 1937, P. P. Babiy, 10° (CU).
CANADA: Quebec, Huntington Co., Covey Hill, Allen Brook, 2 Jul 1966,
D. Koss, 19 (MSU).
Saldula orbiculata (Uhler), 1875
Figures 19 & 21
Discussion--Saldula orbiculata is often brachypterous and in this form
appears very "round". The six pruinose spots on the hemelytra, the dis-
tinctly rounded pronotal margins, and the long hairs on the second antennal
segment and tibiae separate orbiculata from Saldula severini Harris which
lacks the pruinose markings (Fig. 18), has a less rounded pronotum (Fig.
20) and shorter hairs on the antennae and legs.
Habitat preference--Both collections of orbiculata I made in Michigan
were from very damp mossy areas. The Keweenaw County location was
from a sandy margin of a cedar swamp which was covered with scattered
clumps of moss; in Barry County specimens were taken from the Sphagnum
mat near the edge of a bog lake. Wet areas of a permanent nature such as
swamps, marshes, and bogs are required to support this species. The
habits of orbiculata are much like those of Saldula separata (Uhler) in that
it is seclusive and not very mobile. It was collected in association with
Salda bouchervillei (Provancher), Salda obscura (Provancher) and
Micracanthia ripula.
Distribution--The actual identity and distribution of orbiculata is rather
confused. This is primarily a northeastern species, ranging west to the
Rocky Mountains. Records are also known from Mexico (Champion, 1901).
Records from Nevada (Chapman, 1962) and California (Uhler, 1875) are
possibly misidentified specimens of severini. Further collecting will be
necessary to clarify this point.
Specimens examined--MICHIGAN: Barry Co., Purdy Bog, TIN R9W
Sec 36, 2 Aug 1966, T. Schuh, 29; Berrien Co., E. K. Warren Preserve,
Warren Woods, 29 June 1919, R. F. Hussey, 19 (UMMZ); Berrien Co., New
Buffalo, 30 June 1919, R. F. Hussey, 10° (UMMZ); Cheboygan Co., 13 Jul
1949, H. B. Hungerford, 12 (UK); Keweenaw Co., Little Betsy R., T57N
R29W Sec 29, 10 Jul 1966, T. Schuh, F. W. Stehr, 19; Livingston Co.,
E. S. George Reserve, 22 July 1951, I. J. Cantrall, 1c’ (UMMZ); Mecosta
Co., 27 June 1942, R. R. Dreisbach, 1h (UMMZ). COLORADO: Ute Pass,
9 Aug 1925, G. O. Wiley, 12 (UM). ILLINOIS: Galena, 5 Jul 1917, 19
(INHS); Havana, Quiver L., 9 Aug 1895, 19 (INHS); Quiver L., 14 Sept
1894, 12(INHS); Quiver L., 12 Oct 1894, 10° (INHS); Quiver L., Ilinois
R., 10 Nov 1894, 10° (INHS); Lake Forest (beach drift), 13 Jun 1906, J. G.
Needham, 12 (UK). MINNESOTA: LaSueur, crk. 3-1/2 mi. SE, 13 Sept
1923, W. E. Hoffman, 10° (UM); Minneapolis, Glenwood Pk., 9 Jun 1924,
G. O. Wiley, 10°, 12 (UM); Nicollet Co., Barney Fay Ravine, 19 Sept 1923,
W. E. Hoffman, 150’, 172 (UM); Olmsted Co., C. N. Ainslie, 12 (UM).
OHIO: Ironton, 26 May 1899, 1c, 192 (OSU).
20 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Saldula "pallipes-palustris"
Discussion--This species complex has confused the systematic heterop-
terists for some time, and many synonyms exist because of the extreme
variability of the "species". Wagner (1950; reprinted in Usinger, 1956)
gives a eunomic series of the European forms of Saldula pallipes (Fabricius),
1794, and Saldula palustris (Douglas), 1874. The use of the eunomic wing
pattern is fairly reliable for separation of the species in Europe but as yet
has not proved successful in North America. Cobben (1960b; also see 1960a
for discussion of genitalia in separation of European forms) discusses the
occurrence of “palustris" in the Netherlands Antilles, stating that although
the hairs on the processus sensualis of the parameres are short, which
would separate it from pallipes in Europe which has long hairs on the pro-
cessus sensualis, the antillean specimens do not conform to the range of
variation of palustris in Europe. Saldula palustris is primarily halophilous
in Europe which is not the case with most of the specimens designated as
palustris from North America. The parameres of those specimens I have
examined from the Great Lakes region, which appear fromthe wing pattern
to be palustris, are identical with parameres of specimens which appear to
be pallipes. Saldula ablusa closely resembles the "pallipes-palustris" group
(see discussion of ablusa). Differences in the parameres of "pallipes-
palustris" and ablusa exist but their examination is not usually necessary
for separation of the species.
Habitat preference--This form is ubiquitous, and has been collected
throughout the Great Lakes region, and most of the rest of North America
except for the seacoasts. Open stream and pond margins are the most com-
mon habitat and the general collector will usually encounter more represen-
tatives of this group than of all other species combined.
Distribution--I have examined over 1200 specimens from the Great Lakes
region.
Saldula saltatoria (Linnaeus), 1758
Figure 13
Discussion--Appropriately named for its jumping habits, Saldula
saltatoria is one of the more easily recognized species in the genus. The
typical body form is robust, being distinctly broadest at a point near the
apex of the scutellum. The hemelytra are uniformly darkly pigmented; the
eyespot is always distinct, being pruinose, particularly medially and poste-
riorly. Saldula orbiculata is the only other species which has pruinose
markings. The inner apical region of the endocorium also exhibits distinct
pruinose markings. The dorsal vestiture is neat, consisting of short,
golden pubescence on the pronotum and scutellum and some scattered areas
on the hemelytra. Saldula saltatoria most closely resembles orbiculata and
severini but lacks the long, shaggy dorsal vestiture. The European species
Saldula c-album (Fieber) has been recorded from North America, but is
probably an incorrectly identified form of saltatoria.
Habitat preference--Saldula saltatoria is generally found indamp, shaded
areas along ponds and streams. The ground is usually covered with litter
and is seldom exposed to direct sunlight. When disturbed saltatoria moves
very quickly. The dark coloration makes observation difficult in the shaded
habitat.
Schuh: The Shore Bugs of the Great Lakes Region 21
Distribution--Described from Europe, saltatoria is holarctic. Itis
found over most of North America and has been collected as far south as
Guatemala (Champion, 1901). I have examined 35 specimens from the Lower
Peninsula of Michigan collected between April 21 and August 17, and 23 speci-
mens from Illinois, Indiana, Minnesota, New York, and Ontario collected
between March 30 and November 14. I have also seen specimens from the
northwestern United States.
Saldula separata (Uhler), 1878
Figures 22, 23
Discussion--Saldula separata is one of the few species in the genus which
exhibits brachyptery. Of the 17 specimens I collected in Michigan, 6 were
macropterous (40%0’, 229) and 9 were brachypterous (2cc", 799). Figures 22
and 23 show the difference between brachypterous and macropterous forms.
Figure 22 shows a wing with an abberant cell development in the membrane,
a fairly common occurrence in the family. The hairy dorsum and eyes as
well as the distinctive pigmentation of the hemelytra make this species easy
to separate from other members of the genus. Saldula illinoisensis Drake
is a synonym of separata (Polhemus, in press).
Habitat preference--This boreal species has a specialized habitat which
makes collection and observation difficult. It is found in vegetation choked
areas where the ground is covered with mosses and similar low growing
vegetation. The substrate is usually damp to almost dry, with a high or-
ganic content. Saldula separata lies close to the ground and does not fly,
but crawls and jumps short distances. This species was collected in assoc-
iation with Salda obscura.
Distribution--Saldula separata is nearctic and boreal. Scattered records
are known for most of northeastern North America, the range extending from
the Atlantic coast into the Rocky Mountains, and as far south as Colorado.
The southern limit of the eastern part of the range appears to be roughly
southern Illinois.
Specimens examined--MICHIGAN: Charlevoix Co., Dear Crk., near
East Jordan, T32N RTW Sec 8, 22 Jul 1966, T. Schuh, E. Evans, 1c, 39;
Douglas L., 23 Jul 1927, H. B. Hungerford, 10 (UK); Grand Traverse Co.,
Marion Island, 25 July 1923, T. H. Hubbell, 20°, 42 (UMMZ); Keweenaw Co.,
Montreal R., T58N R30W Sec 14, 10 Jul 1966, T. Schuh, F. W. Stehr, 29;
Keweenaw Co., T57N R30W Sec 36, 10 Jul 1966, T. Schuh, F. W. Stehr,
1c", 29; Manistee Co., 7-5-52, R. R. Dreisbach, 12 (MSU); Ontonogan Co.,
Sooty Creek, T50N R44W Sec 24, 11 Jul 1966, T. Schuh, 40°, 49. ILLINOIS:
N. Ill., Andreas Bolter Coll., 12 (INHS). MINNESOTA: St. Paul, 1 Jul
1923, G. O. Wiley, 22 (UM). NEW YORK: Adirondack Mts., Mt. Marcy,
9 Jul 1918, C. R. Crosby, 19 (CU). CANADA: Ottawa, Torre-Bueno Coll.,
1c’, 12 (UK); Ontario, Sanford, 1906, 192 (CU).
Saldula severini Harris, 1943
| Figures 18 & 20
Discussion--See discussion under Saldula orbiculata (Uhler).
22 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Habitat preference--The habitat of S. severini is similar to that of
orbiculata.
Distribution--Collection records indicate that severini is confined pri-
marily to the Rocky Mountains and the western part of the Great Plains.
Harris (1943) records specimens from Waubay, South Dakota and Palo Alto
County, Iowa.
Specimens examined--SOUTH DAKOTA: Custer St. Pk., 20 June 1955,
R. L. Fischer, R. W. Hodges, 20°, 22 (MSU).
TRIBE SALDINI
Salda Fabricius
The genus Salda is holarctic with most of the species showing a boreal
distribution. Reuter (1912) established the genus Lampracanthia to include
a single species, crassicornis Uhler, and the genus Teloleuca to include a
number of species, two of which occur in North America, pellucens
(Fabricius) and bifasciata (Thompson). Cobben (1959) revised the status
of the genus Salda and included in it Lampracanthia and Teloleuca as sub-
genera. This change of status was based on a study of the genitalia and
other morphological characters, and consideration of the distribution and
biology of the species concerned. Representatives of all three subgenera
occur in the Great Lakes region, although apparent specificity of habitat
makes them very difficult to collect.
Salda littoralis (Linnaeus), a European and Asian species, has been
recorded from eastern North America in the literature. There is some
question as to the validity of the name for this species in North America,
but I have included it in the keys to make separation of the species easier
(see Polhemus, in press for discussion).
Salda (Teloleuca) bifasciata (Thompson) (Fig. 4), a European, Asian,
and western North American species has also been included in the keys
for completeness.
Salda (Salda) anthracina Uhler, 1875
Discussion--Salda anthracina is uncommon in collections. As are most
other members of this genus, anthracina is quite variable morphologically.
It can be distinguished from S. bouchervillei (Provancher) by the companu-
late pronotum, which is only slightly explanate laterally. The antennae are
stout (proportions - I-3, II-2, IV-2) whereas those of bouchervillei are
quite slender (Fig. 30) (proportions - II-5, III-3, IV-3).
Habitat preference--Little is known about the habitat preference of
anthracina because itis souncommon. It apparently lives in situations
similar to those preferred by bouchervillei.
Distribution--Salda anthracina was described from York County,
Pennsylvania (Uhler, 1875). Drake and Hottes (1950) also record it from
New Hampshire and New York. John Polhemus (correspondence) has ex-
amined specimens in the Drake Collection from New Jersey and Tennessee.
I have seen one specimen from Berrien County, Michigan.
Specimens examined--MICHIGAN: Berrien Co., E. K. Warren Preserve,
Warren Woods, 25 June 1919, R. F. Hussey, 19 (UMMZ). OHIO: Sugar
Schuh: The Shore Bugs of the Great Lakes Region 23
Grove, 17 May 1902, Herbert Osborn Collection, 12 (OSU). NEW YORK:
Dryden, Ringwood, 6 Jul 1950, H. Dietrich, 59 (CU).
Salda (Salda) bouchervillei (Provancher), 1872
Figure 30
Discussion--Salda bouchervillei is one of the most common eastern
species in the genus. The dorsum is shining black, with only scattered
indistinct golden pubescence, whereas S. buenoi (McDunnough) has a dull
black dorsum with only slightly shining appressed pubescence. The lateral
pronotal margins are nearly straight and explanate posteriorly, Salda
anthracina has a more companulate pronotum and less explanate lateral
pronotal margins. Brachypterous specimens, which are the most common,
are robust (length 5.5 to 6.5 mm., width 2.8 to 3.3 mm.) and convex dor-
sally; the margins of the hemelytra are evenly rounded and noticeably ex-
planate anteriorly. Macropterous specimens are much more elongate
(length 6.5 to 7.5 mm., width 2.8 to 3.3 mm.), the hemelytra appearing
almost straight-sided. All antennal segments are uniformly colored, and
usually very dark, whereas buenoi has a light second segment. The first
segment has a light stripe on the interior edge; this stripe is sometimes
very indistinct. Salda anthracina has stouter antennae with a light second
segment, and in S. obscura (Provancher) the second antennal segment is
dark with a light stripe dorsally. The legs are nearly uniformly colored,
being almost without dark pigment in males and almost entirely black in
the females.
Habitat preference--Salda bouchervillei inhabits marshy areas with
dense vegetation. It is seclusive, hiding at the base of moss or grass
clumps. When disturbed it darts out, but quickly seeks cover again. Most
specimens are brachypterous and flight is therefore uncommon.
Distribution--Salda bouchervillei has a range similar to that of S.
obscura, but it does not appear to occur in the northwestern United States.
Drake and Hottes (1950) cite records from Alaska, New Mexico, and Ten-
nessee. The New Mexico records are probably incorrectly identified
specimens of Salda buenoi.
Specimens examined--MICHIGAN: Bay Co., 18 Jun 1950, R. R.
Driesbach, 10, 19 (MSU); Berrien Co., E. K. Warren Reserve, Sawyer
Dunes, 24 June 1919, T. H. Hubbell, 19 (UMMZ); Berrien Co., E. K.
Warren Reserve, Warren Woods, 15 July 1920, T. H. Hubbell, 1c’ (UMMZ);
Charlevoix Co., Garden Island, 28 July 1920, S. Moore, 12 (UMMZ);
Charlevoix Co., HighIsland, 14 June 1922, S. Moore, 19 (UMMZ); Charle-
voix Co., High Island, 7 July 1922, S. Moore, 12 (UMMZ); Cheboygan Co.,
18 Jul 1931, H. B. Hungerford, 10 (UK); Cheboygan Co., 24 Jul 1930, H. B.
Hungerford, 12 (UK); Cheboygan Co., Douglas L., 22 June 1925, C. F.
Byers, 12 (UMMZ); Cheboygan Co., Monroe L., on beach, July 1930,
J. Kigis, 1c’, 12 (UMMZ); Cheboygan Co., Mud L. Bog, T38N R3W Sec 7,
24 Jul 1966, T. Schuh, E. Evans, 20°, 192; Emmet Co., Petoskey, T35N
RdoW Sec 34, 23 Jul 1966, T. Schuh, E. Evans, 10°; Keweenaw Co., Little
Betsy R., T57N R29W Sec 29, 10 Jul 1966, T. Schuh, F. W. Stehr, 100,
99; Keweenaw Co., T57N R30W Sec 36, 10 Jul 1966, T. Schuh, F. W. Stehr,
40°, 69; Mackinac Co., Naubinway, 24 June 1921, S. Moore, 19 (UMMZ);
Marquette Co., Peshekee R., T48N R30W Sec 25, 9 Jul 1966, T. Schuh,
24 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
F. W. Stehr, 1c’, 19; Oakland Co., Milford, 22 June 1921, T. H. Hubbell,
1h’ (UMMZ); Ontonogan Co., Union R. at L. Superior, 11 Jul 1966, T. Schuh,
lo’. COLORADO: Sedalia, 18 Jul 1962, J. T. Polhemus, 1c", 19 (TS).
INDIANA: Pine (lake beach), 28 May 1905, W. J. Gerhard, 10°, 2? (FM);
Wolf L., 11 Jun 1911, A. B. Wolcott, 1c (FM). ILLINOIS: Chicago, 5 Jun
1904, W. J. Gerhard, 12 (FM); Chicago, Avalon Park, 15 Jun 1933, W. J.
Gerhard, 19 (FM); Chicago, Dauphin Park, 19 Jun 1920, A. B. Wolcott, 19
(FM); S. Chicago, 7 Jul 1907, W. J. Gerhard, 10°, 19 (FM); South Chicago,
0 Jun 1910, A. B. Wolcott, 12(FM); N. Ill., Andreas Bolter Coll., 49
(INHS); Urbana, 1 Jun 1889, Hart Coll., 22 (INHS). IOWA: Lake Park,
12 Jun 1948, T. Polhemus, 1c (FM). MINNESOTA: Chisago Co., Rus
Beach, 3 Jun 1900, Otto Lugger Coll., 49 (UM); Ottertail Co., Otto Lugger
Coll., 12(UM). NEW YORK: N. Fairhaven, 4-?-21, 12(CU). NORTH
DAKOTA: Devil's L., 23 July 1920, 49 (M); Nelson Co. , Stump L., 24 July
1920, T. H. Hubbell, 19 (UMMZ).
Salda (Salda) buenoi (McDunnough), 1925
Discussion--Salda buenoi and Salda bouchervillei are very similar in size
and shape. The dorsum of buenoi is a dull black with appressed, slightly
shining pubescence, whereas bouchervillei is highly polished and almost
devoid of pubescence (Fig. 30). Salda littoralis can be separated from buenoi
by its dense, shining, golden pubescence.
Habitat preference--Salda buenoi is found in damp areas with sedge and
grass cover. It is often in more exposed areas than bouchervillei, on sandy
or muddy margins of streams and ponds, in a habitat similar to S. lugubris
(Say).
Distribution--Salda buenoi has been recorded from Washington, Oregon,
California, Idaho, Nevada, Montana, Colorado, Utah, Nebraska, North
Dakota, South Dakota, Iowa, Illinois, Wisconsin, Michigan, Massachusetts,
New Hampshire, British Columbia, Alberta, Saskatchewan, Ontario (Drake
and Hottes, 1950) and Newfoundland (Lindberg, 1958). Polhemus Leer res:
pondence) has seen specimens from Arizona.
Specimens examined--COLORADO: Hygiene, 12 May 1963, John T.
Polhemus, 10°, 19 (TS). UTAH: Uba Dam, Sevier R., B. Moore, 19
(UMMZ).
Salda (Salda) lugubris (Say), 1831
Discussion--The generic position of Salda lugubris has been much dis-
puted. Cobben (1959) places it in Salda, rather than Saldula where it was
previously placed, because of the cell structure of the membrane. The
inner cell of the membrane is produced no more than one-fourth of its total
length beyond the base of the second cell, whereas the rest of the members
of the genus have the inner cell produced about one-third of the total length.
Drake and Hottes (1950) mention that both long- and short-winged forms are
‘common, but I have never seen a brachypterous specimen. Salda lugubris
most closely resembles the macropterous form of buenoi, but can be sepa-
rated from it by the difference in the cells of the membrane. The pubescence
of lugubris is more dense and golden than that of buenoi and Saldula nigrita.
Salda lugubris is much more robust than nigrita. Saldula major (Uhler) is a
synonym of lugubris.
Schuh: The Shore Bugs of the Great Lakes Region 25
Habitat preference--The typical habitat is a sandy shore of a pond or
stream with some emergent vegetation, usually sedges or grasses. Salda
lugubris was collected on the shore of Lake Superior in Baraga County,
Michigan, hiding in the matted sedges on the sandy lake shore. The species
runs and flies very rapidly when disturbed. Wiley (1922) reared lugubris
and gives a brief life history of it.
Distribution--Salda lugubris has a rather atypical distribution for the
genus. Drake and Hottes (1950) record this primarily eastern North American
species from as far west as British Columbia, Utah and New Mexico. The
southern range of lugubris is much greater than any other member of the
genus. I have examined one specimen from Florida. I have examined 38
specimens from Michigan collected between June 12 and Aug 12, and 56 speci-
mens from Florida, Illinois, Indiana, Massachusetts, Minnesota, Montana,
New York, Wisconsin and Ontario collected between April 9 and Oct 2.
Salda (Salda) obscura (Provancher), 1872
Discussion--Salda obscura is one of the smallest members of this sub-
genus (length 3.8 to 5.6 mm., width1.7 to 2.6 mm.). The dorsum is shining
black and almost devoid of pubescence, which separates it from littoralis
which has dense, shining, golden pubescence. The lateral margins of the
pronotum are very straight, and the lateral margins of the hemelytra only
slightly rounded, giving the appearance of being widened posteriorly. All
of the specimens I have examined have been brachypterous. The antennae
are covered with short, golden, subappressed pubescence and some longer
erect dark hairs. The first and second antennal segments are light dorsally
and black ventrally; antennal segments three and four are entirely black;
bouchervillei has a completely dark second segment and in littoralis the
second segment is entirely light. Antennal proportions are: I-20; II-14;
IV-14. The legs are mostly light, the femora have a dark "saddle" apically;
-The amount of dark pigmentation is, however, quite variable.
Habitat preference--Collections from Michigan have shown obscura to
live on fairly dry ground, with some vegetation. In Keweenaw County,
Michigan it was collected on sandy areas at the edge of a cedar swamp; both
obscura and bouchervillei were present in this habitat. The substrate was
covered with scattered clumps of mosses with sundew plants growing in
them; the saldids were found hiding in the moss clumps and would dart out
periodically. Locations where I have collected obscura in Oregon were damp,
sandy slopes near streams. The saldids, often present in large numbers,
were found running among the low,. scattered vegetation. Lindberg (1958)
notes collecting obscura on marshes with Spagnum-Carex-Myrica and Carex-
Juncus, on riverbanks, and on the seashore. Saldula separata and Micra-
canthia ripula were collected in association with obscura.
Distribution--Salda obscura, like most other members of the genus, is
a true boreal form, and has been collected in the mountains in British
Columbia, Washington, Oregon, Nevada, Colorado (Chapman, 1962), British
Columbia and Washington. Lindberg cites collections from Newfoundland,
— Quebec, Alberta and Alaska. All records from Michigan are from areas near
the southern shore of Lake Superior.
Specimens examined--MICHIGAN: Keweenaw Co., T57N R30W Sec 36,
10 Jul 1966, Toby Schuh, F. W. Stehr, 10°; Keweenaw Co., Little Betsy R.,
TO7N R29W Sec 29, 10 Jul 1966, Toby Schuh, F. W. Stehr, 10’, 29;
26 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Ontonogan Co., Sooty Crk., T50N R44W Sec 24, 11 Jul 1966, Toby Schuh,
1c’; Ontonogan Co., Mineral R. at L. Superior, 11 Jul 1966, Toby Schuh,
12. OREGON: Umatilla Co., 5 mi. W. Tollgate, el. 4000 ft., 11 Jul 1964,
J. D. Lattin, Toby Schuh (TS); Union Co., 6 mi., E. Tollgate, el. 4900 ft.,
12 Jul 1964, J. D. Lattin, Toby Schuh (TS).
Salda (Lampracanthia) crassicornis Uhler, 1875
Discussion--This species is easily distinguished from other members of
the genus by its small size (length 3.0 to 3.5 mm.) shining black dorsum and
sparse erect vestiture. Most specimens are brachypterous but macropterous
forms have been taken (Slater, 1955).
Habitat preference--Slater (1955) notes tedbiy Salda crassicornis ina
marshy area adjacent to the Niobrara River, at Valentine, Nebraska. Many
specimens were found by Slater on matted marsh grass and sedge in water
from ankle to knee depth. Lindberg (1958) collected crassicornis in New-
foundland on humid marshes, swamps with Carex and Myrica and on the
shores of spagnum pools.
Distribution--Although crassicornis has not been collected in Michigan
it should occur here. Records from the literature and available collections
indicate a boreal distribution. Blatchley (1926) collected four specimens on
the shore of Lake Tippacanoe, Koscinsko County, Indiana, between June 7
and 9.
Specimens examined--NEW YORK: Dryden, Ringwood, 6 Jul 1950,
H. Dietrich, 2% (CU).
Salda (Teloleuca) pellucens (Fabricius), 1779
Figure 3
Discussion--The holarctic subgenus Teloleuca has two species which
occur in North America, pellucens (Fabricius) and bifasciata (Thompson).
The dorsal facies more closely resembles that of Saldula than the typical
Salda (Figs. 3 & 4). Salda pellucens can be easily separated from bifasciata
by the presence of a light quadrangular area at the lateral apex.of the endo-
corium (Fig. 3), whereas there is only a small white spot at this location
in bifasciata (Fig. 4).
Habitat preference--Salda pellucens is most often collected along gravel
or sand stream margins, but may occur some distance from permanent
water. It seems to be very positively phototrophic and does not remain in
the open unless there is bright sunlight.
Distribution--This species is strictly boreal and occurs across most of
Canada and the northern United States as well as in Europe and Russia
(Drake, 1952). Records are known from Essex Co., New York (Blatchley,
1926; cited as Chartoscirta (Chartolampra) cursitans Bueno), Montana,
Colorado and northern Wisconsin. I have examined specimens collected
by J. T. Polhemus at Weston Pass, Colorado. It should occur in Michigan.
Harold Chapman collected a specimen in Lakewood, Wisconsin.
Schuh: The Shore Bugs of the Great Lakes Region 27
LITERATURE CITED
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Publishing Co., Indianapolis, Indiana, 1116 pp.
Champion, G. C. 1897-1901. Insecta. Rhynchota. Hemiptera-Heteroptera.
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Chapman, H. C. 1962. The Saldidae of Nevada. Pan-Pac. Ent. 38: 147-
159.
Cobben, R. H. 1957. Beitrag zur Kenntnis der Uferwanzen. Ent. Ber.
17: 245-57.
1959. Notes on the classification of Saldidae with the description
of a new species from Spain. Zool. Mededel. 36: 303-16.
. 1960a. Monographische Bearbeitung der Saldiden Europas.
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Drake, C. J. 1949. Some American Saldidae. Psyche .56: 187-193.
. 1950. Concerning North American Saldidae. Bull. Brooklyn
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. 1952. Alaskan Saldidae. Proc. Ent. Soc. Washington. 54:
145-48.
. 1958. Anundescribed shorebug from Manitoba. Bull. S. Calif.
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Drake, C. J. and H. C. Chapman. 1952. A new species of Micracanthia
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. 1953. An undescribed saldid from the Gulf States. Bull.
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.. 1958. The subfamily Saldoidinae. Ann. Ent. Soc. Amer.
D1: 480-85. ,
Drake, C. J. and L. Hoberlandt. 1951. Catalogue of genera and species
of Saldidae. Acta Ent. Mus. Nat. Prague. 26, No. 376: 12 pp.
Drake, C. J. and F. C. Hottes. 1950. Saldidae of the Americas. Great
Basin Nat. 10: 51-61.
1954. Synonymic data and a description of a new saldid. Occ.
Pap. Mus. Zool., Univ. Michigan. No. 553: 5 pp.
Ekblom, T. 1926. Morphological and biological studies of the Swedish
families of Hemiptera-Heteroptera. PartI. Zool. Bidr. 10: 31-180.
Fabricius, J. C. 1779. Reise nach Norwegen mit Bemerkungen aus der
Naturhistorie und Oekonomie. Hamburg, Bohn, 388 pp.
. 1794. Entomologica systemica emendata et aucta. Hafniae.
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Froeschner, R. C. 1962. Contributions to a synopsis of the Hemiptera
of Missouri, Part V. American Mid. Nat. 67: 208-40.
Harris, H. M. 1937. Contributions to the South Dakota List of Hemiptera.
Iowa State College J. Sci. 11: 169-76.
1943. Additions to the South Dakota List of Hemiptera. J.
Kansas Ent. Soc. 16: 150-53.
28 Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
Lindberg, H. 1958. Hemiptera-Heteroptera from Newfoundland collected
by Swedish-Finnish Expedition of 1949 and 1951. Acta Zool. Fenn.
96: 1-25.
Linnaeus, C. 1758. Systema Naturae. 10th Ed. Vol. 1: 448.
McDunnough, J. (Description of Salda buenoi). Canad. Ent. 57: 259-60.
Osborn, H. 1901. A new genus, including two new species of Saldidae.
Canad. Ent. 33: 181-82.
Parshley, H. M. 1921. A report on some Hemiptera from British Columbia.
18 (Systematic Series): 13-24.
Polhemus, J. T. 1967. In Press.
Provancher, L. 1872. Descriptions de plusieurs Hemipteres nouveaux.
Nat. Canad. 4: 106-07.
Reuter, O. M. 1912. Zur generishcen Teilung der Paldarktischen und
nearktischen Acanthiaden. Ofv. Finska Vet-Soc. Fohr. 54: 1-24.
Say, Thomas. 1831. Descriptions of new species of heteropterous Hemiptera
of North America. New Harmony, Indiana.
Scudder, G. G. E. 1961. Some Heteroptera new to British Columbia.
Proc. Ent. Soc. British Columbia. 58: 26-29.
Slater, J. A. 1955. The macropterous form of Lampracanthia crassicornis
(Uhler). J. Kansas Ent. Soc. 28: 107-09.
Uhler, P. R. 1875. Report on the insects collected by P. R. Uhler during
the explorations of 1875. Bull. U.S. Geol. Geog. Surv. Terr. 1:
355-475.
. 1877. List of Hemiptera of the region west of the Mississippi
River. Bull. U.S. Geol. Geog. Surv. Terr. 3: 331-334.
. 1878. Notices of the Hemiptera-Heteroptera in the collection
of the late T. W. Harris, M.D. Proc. Boston Society Natural History.
19: 432.
Usinger, R. L., Editor. 1956. Aquatic Insects of California. University
of California Press, Berkeley, 508 pp.
Wagner, E. 1950. Notes on Saldidae. Acta Ent. Mus. Nat. Prague.
26, No. 371: 4 pp. :
Wiley, G. O. 1922. Life history notes on two species of Saldidae found in
Kansas. Kansas Univ. Sci. Bull. 14: 301-11.
Zetterstedt, J. W. 1840. Insecta Lapponica descripta. Lipsiae. col. 268.
Schuh: The Shore Bugs of the Great Lakes Region 29
measurement of, vertex
and eye
e+ vertex
ie
60. 9ce [lus
<—_—_—__—_—_ — compoun
—__—__—_ pronotal cotlar
—<—__—_—_—_ anterior lobe
——————_—_—_—— pronotum
posterior lobe
el daitil tanita OTA VSS
/ gees
a
~-——$_—exocorium
Viv
Hava pas Ae
1 membrane
Fig. 1.--General morphological details. Fig. 2.--Left hemelytron, Pentacora
ligata (Say). Fig. 3.--Left hemelytron, Salda (Teloleuca) pellucens (Fabricius)
o& (Colorado, Weston Pass). Fig. 4.--Left hemelytron, Salda (Teloleuca)
bifasciata (Thompson) & (Washington, Mt. Ranier National Park).
30
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
5. --General dorsal habitus, Saldoida turbaria new species ?
(Michigan, Barry County, Purdy Bog)
6. --Lateral view, head and thorax, Saldoida turbaria 2
7. --Ventral view, apex of abdomen, Saldoida turbaria ¢
8.--Left hemelytron, Micracanthia humilis (Say) ?
(Michigan, Gratiot County, Maple River at Banister)
9.--Left hemelytron, Micracanthia humilis (Say)
10.
11.
12.
13.
14.
15.
16.
1%,
(Michigan, Gratiot County, Maple River at Banister)
--Left hemelytron, Micracanthia humilis (Say) 2
(Michigan, Gratiot County, Maple River at Banister)
--Left hemelytron, Micracanthia fennica Reuter ?
(Colorado, Hartsel
--Left hemelytron, Micracanthia quadrimaculata (Champion) “
(Oregon, Baker County, 4 miles W. Baker, North Powder
River)
--Left hemelytron, Saldula saltatoria (Linnaeus) ©
(Michigan, Shiawassee County, Rose Lake Exp. Station)
--Left hemelytron, Saldula ablusa Drake and Hottes ?
(Michigan, Cheboygan County, Douglas Lake)
--Left hemelytron, Saldula ablusa Drake and Hottes 9
(Michigan, Emmet County)
--Left hemelytron, Saldula ablusa Drake and Hottes &
(Michigan, Alger County, Kingston Lake)
--Left hemelytron, Saldula ablusa Drake and Hottes ?
(Michigan, Menominee County, T34N R25W Section 20)
32
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
18
19
20
21
22
23
24
29
26
27
Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
.--Left hemelytron, Saldula severini Harris &
(South Dakota, Custer State Park)
.--Left hemelytron, Saldula orbiculata (Uhler) ?
(Michigan, Keweenaw County, Little Betsy River)
.--Pronotum, Saldula severini (Harris) &
(South Dakota, Custer State Park)
.--Pronotum, Saldula orbiculata (Uhler) ¢
(Michigan, Keweenaw County, Little Betsy River)
.--Left hemelytron, Saldula separata (Uhler) 2? macropterous
(Michigan, Keweenaw County, Montreal River,
T58N R30W Section 14)
.--Left hemelytron, Saldula separata (Uhler) ¢ brachypterous
(Michigan, Ontonogan County, Sooty Creek,
T50N R44W Section 24)
.--Left hemelytron, Saldula opacula (Zetterstedt) &
(Michigan, Kalkaska County, Desmond Creek)
.--Left hemelytron, Saldula nigrita Parshley “
(Michigan, Baraga County, Otter River)
.--Left hemelytron, Saldula confluenta (Say)
(Michigan, Emmet County, Maple River at Brutus Road)
.--Left hemelytron, Saldula comatula (Parshley) &
(California, Siskiyou County, Medicine Lake)
“(PT UOTOEg MOSH NBGL ‘1eaTY TeerjUOW ‘4QunoD meusemey ‘ueSTYyoTI) . ayeaq e[NdIa eIIpUBOeIOTIN-- °6Z “Sy
*(peoy snjynig ye JaATY [den ‘AjunoD JowWWy ‘UBSTYSTIA) ,.o ueudeyD pue oyeiq euUEpIIOT] BIUJUBOeIOI\-- "97 “Sly
Contr. Amer. Ent. Inst., vol. 2, no. 2, 1967
34
Schuh: The Shore Bugs of the Great Lakes Region 39
Fig. 30. --Salda (Salda) bouchervillei (Provancher) ? brachypterous (Michigan,
Keweenaw County, Little Betsy River).
yey ‘i i
Pein
DL
Yl, /
As|
Enl,
Contributions
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American Entomological Institute
Volume 2, Number 3, 1967 ©
~~
THE CHAETOTAXY OF NORTH AMERICAN
LEPIDOCYRTUS S. STR., (COLLEMBOLA: ENTOMOBRYIDAE)
By Richard J. Snider
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Contributions
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Volume 2, Number 3, 1967
"Vr
THE CHAETOTAXY OF NORTH AMERICAN
LEPIDOCYRTUS S. STR., (COLLEMBOLA: ENTOMOBRYIDAE)
By Richard J. Snider
TABLE OF CONTENTS
PR Ga Ba te ey as ba es he es i a a ew a he eee
CSE QOS OR ESN Soa. ig 1 GENREUAUIRI Ets MADRS GESRI PRR atin (een ee ROO CRU Ne ee aa
A Es Pe Be ee es co acca cen dh ade, dans aed ek we ed we ee ie ew
COR ee ida ry se ite ahigial ae ata Men dia blob. Ware Baw ee 0
Boo (Phe SINC Poche Fat Or i ae CNG ek wa hea a a we eee ees
Il. The Microchaetae (Accessories) Associated with
the Lateral Lasiotrichia of Abdomen III...............
III. The Microchaetae (Accessories) Associated with
the Anterior Lasiotrichia of Abdomen IV..............
CHAETOTAXY OF NORTH AMERICAN SPECIES..........ceeecccces
PE A Eat arias alg aaa hae ta ae a ar Le ae A, Wee) cat as ab, Suse gece ta nk eae wa, as eee
i ee he es eg ths bin uiaiiws ich Gow iene Mabie pica oun ase, Romane we
Pe ete NS EHOW a VAN ee eRe ee MUO W NT Oe Wee ee ee a oe
THE CHAETOTAXY OF NORTH AMERICAN
*
LEPIDOCYRTUS S. STR., (COLLEMBOLA: ENTOMOBRYIDAE)
By
Richard J. Snider
Dept. of Entomology, Michigan State University, East Lansing, Michigan
ACKNOWLEDGMENTS
I wish to thank the following individuals: Dr. Roland Fischer, Department
of Entomology, and Dr. T. Wayne Porter, Department of Zoology of Michigan
State University for their kind cooperation during the writing of this manu-
script. Special thanks are extended to: Dr. James Butcher, Department of
Entomology, for consultation and encouragement; Dr. Kenneth Christiansen,
Grinnell College, for aid in limiting and outlining the topic; Dr. Hermann
Gisin, Museum d' Histoire Naturelle de Geneve, for his criticism and addi-
tional suggestions; Dr. Harlow B. Mills, Chief (retired), the Illinois Natural
History Survey, for many years of aid and understanding.
INTRODUCTION
The genus Lepidocyrtus Bourlet is a member of the tribe Lepidocyrtini
which includes: Pseudosinella, Seira, and Heteromurus.
The genera listed above are very closely related and are often separated
on the basis of presence or absence of eyes, falcate or toothed mucro, and
whether or not the 4th antennal segment is simple or annulate. In former
works members of the Lepidocyrtini were sometimes included under the
genus Lepidocyrtus. Recognition of the North American genera as separate
entities was done by Mills (1933). In the United States the genus Lepidocyrtus
is recognized by the following criteria: eyes 8 on each side of the head; body
with finely striated scales; lateral tooth on either side of the unguis, basal
inner teeth not winglike; well developed tenent hair present; mesonotum
projects over the reduced pronotum, obscuring it; ventral surface of the
dentes with scales.
The problems involved in making generic and specific determinations of
some members of the order Collembola concern finding reliable morphological
_ key characters to separate them one from the other. The small size of Col-
lembola forced early workers to restrict their observations to limited
ok
This research was partially supported by an N.I.H. grant, no. CC00246.
2 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
anatomical differences and color patterns. When Sir John Lubbock (1873)
published his great monograph on the Apterygota, a more organized and ex-
acting approach to the study of Collembola began. He observed and recorded,
both in text and figures, minute structures ignored by earlier taxonomists.
Detailed comparative systematic work on the order was initiated at the be-
ginning of the 20th century with the work of Carl Bérner (1901 and 1906) and
until recently has not changed significantly in character.
The first advances in chaetotaxy began with Bonet (1945), who described
the chaetotaxy of the head of some species of Hypogastrura. Delamare-
Debouttville (1951) surveyed the known information on macrochaetae and
trichobothria associated with soil-type Collembola and related these struc-
tures to phylogeny. Yosii (1956) described the basic chaetotaxy of the
Hypogastruridae and distinguished each seta of the head and body segments.
Cassagnau (1959) discussed the distribution of macrochaetae on the tergites
of Tetracanthella. He stated that the position of the macrochaetae was of
great systematic importance. Using chaetotaxic patterns he was able to
divide Tetracanthella into three chaetotaxic types. In his revision of the
genus Entomobrya, Christiansen (1958) used chaetotaxy to a limited extent.
He divided the body setae into five types based on shape and lengths, but made
no use of gross body setal patterns except on the male genital plate. Yosii
(1959) revised the genus Seira and utilized the distribution of the macrochaetae
on the tergites. Gisin (1960) used chaetotaxy of various groups to advantage
in his discussion of species and included setal pattern drawings for his new
species.
As Yosii has pointed out in an earlier paper (1956), the use of chaetotaxy
is an important taxonomic tool. Yosii (1960) refined his work on Hypogastrura
and attempted to define the evolution of the selected species he worked with
and to show affinities. Once again using chaetotaxy, Yosii (1961) surveyed the
Collembola, and related the family groups phylogenetically.
Gisin (1961) in reworking the collection of Carl Bérner elaborated the
species descriptions using chaetotaxy. Gisin and DaGama (1962) compared
the chaetotaxy of three species of Seira. They found that the setae showed
differences in relative distances from each other. As chaetotaxic technique
developed, authors put it to use in difficult situations. Cassagnau (1964) made
a comparative study of the dorsal chaetotaxy of Hypogastrura and was able to
group Closely allied species by pattern type. Massoud (1964) used a chaeto-
taxic table in describing a new species. Christiansen (1964) used position
of the macrochaetae for specific identification. He also mentioned that the
microsetae associated with the trichobothria are constant in position.
Murphy (1966) discussed the taxonomy and bionomics of Sphaeridia,
utilizing the dorsal head and body chaetotaxy. He assigned a notation system
to the setae which was followed in his descriptions of species. Betsch and
Cassagnau (1966) described the evolution and form of the chaetotaxy of the
abdominal papillae of Arrhopalites from the juvenile to the adult stage.
Christiansen (1966), in a revision of Arrhopalites, systematically used chaeto-
taxy to characterize species.
The understanding of the chaetotaxy for the genus Lepidocyrtus has been
largely due to the efforts of one man, Dr. Hermann Gisin. In a paper dealing
with the Lepidocyrtini, Gisin (1963) discussed the pseudopores and chaetotaxy
of this tribe, and illustrated the chaetotaxic pattern for Pseudosinella, a
genus closely allied to Lepidocyrtus. In this paper he outlined a basic chae-
totaxic map analogous to the setal maps used in describing lepidopterous
larvae, and set down a notation system for the tribe Lepidocyrtini.
Snider: The Chaetotaxy of Lepidocyrtus S. Str. 3
Gisin (1964a) presented a partial revision of the genus Lepidocyrtus s. str.
in which he was able to show differences between species. This first article |
was concerned with seven species. Using only the mid-dorsal setae, he man-
aged to develop four categorical chaetotaxic groups within the genus for Euro-
pean species: 1) Lepidocyrtus curvicollis Bourlet, Lepidocyrtus violaceous
Geoffroy and Lepidocyrtus instratus Handschin; 2) Lepidocyrtus languginosus
Gmelin and Lepidocyrtus cyaneus Tullberg; 3) Lepidocyrtus paradoxus Uzel;
4) Lepidocyrtus fimetarius Gisin.
Previously L. violaceous Geoffroy and L. cyaneus Tullberg had been dif-
ficult to separate. He noted that they could be distinguished with chaetotaxic
characters. This discovery eventually led to other usable morphological dif-
ferences in separating these two very similar species. Dr. Gisin showed that
the dorsal chaetotaxy of abdominal segment IV differed between L. cyaneus
and L. violaceous on the microchaetal level. Gisin (1964b) later developed
more refined techniques for their separation. His studies of the genus
Lepidocyrtus led to the use of the taxonomic characteristics of fine structures
associated with the macrochaetae which had heretofore never been considered.
Gisin was able to separate L. lignorum Fabricius from L. curvicollis Bourlet,
a species that had up to this time masked lignorum on general morphological
characters.
Gisin (1965) separated Lepidocyrtus pallidus Reuter from L. cyaneus
Tullberg using chaetotaxy. At the same time he distinguished Lepidocyrtus
serbicus Denis from L. pallidus Reuter and described a new species,
Lepidocyrtus felxicollis Gisin, a species closely related to L. curvicollis.
Hale (1966) made use of the precepts laid down by Gisin and studied L.
lignorum intensively, demonstrating that there is some variation in macro-
chaetal size between individuals of the same species.
Two of the species described as new in this paper were observed by
Dr. Justus Watson Folsom in the late nineteen twenties. Dr. Folsom made
preliminary observations of the species described herein, Lepidocyrtus
millsi n. sp. and Lepidocyrtus floridensis n. sp., but never published his
results. Through Dr. Harlow Mills the efforts of Dr. Folsom were made
available. I have made use of Dr. Folsom's habitus drawings where appli-
cable and have given him credit. Death prevented completion of work on the
genus Lepidocyrtus by Dr. Folsom. It is to his credit that work 30 years
old has stood the test of time and is still valid.
MATERIALS AND METHODS
The specimens used in this study were obtained, where possible, from
the collections in the Entomology Museum, Michigan State University. This
collection was augmented by specimens of Lepidocyrtus on loan from: the
Illinois Natural History Survey; the United States National Museum; the
University of Michigan Museum; the collection of Dr. Kenneth Christiansen
of Grinnell College, Iowa; the collection of Dr. H. Goto, Imperial College,
London, England; and the Museum of Natural History, Geneva, Switzerland
through Dr. Hermann Gisin. Without the cooperation of these sources this
study would not have been possible.
All specimens which have been examined were taken from alcohol-
preserved collections, as previously made slide mounts were not usually
of the nature which allowed good microscopic work. In my experience, 95%
Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
ethanol is the best all around preservative. It tends to make the specimens
stiff, but the color and integrity of the exoskeleton remain intact much better
than in other preservative solutions which have been tried.
To make observations of the chaetotaxy in the genus Lepidocyrtus, it was
necessary that specimens be placed on microscope slides. Specimens of all
species described in this paper were prepared in the following manner:
iL
2.
8.
9.
The individuals were placed in 95% ethanol if not previously preserved
in the same.
They were decapitated and the appendages removed, including the collo-
phore. The head was placed aside for separate mounting as it is easily
lost in a watch glass of alcohol.
With very fine pins mounted in match stick handles, a cut was made along
the mid-ventral line from prothorax to anus. The specimen was held by
the furcula while the cutting was done.
The preparation was then placed into a 5.25% solution of sodium hypo-
chlorite. The clearing of the specimen must be closely observed since
it proceeds very rapidly (3-5 minutes). Clearing in such a manner insures
that all body contents are dissolved with nothing remaining except the
exoskeleton.
The cleared exoskeleton was rinsed twice with distilled water (5 minutes
each). This removes the sodium hypochlorite.
The specimen was then placed on a microscope slide dorsal side up ina
drop of water, and the slide was tilted at an angle so the water drained
from beneath the specimen. Excess water was removed with blotting
paper.
A drop of CMC-10* was placed next to the wet preparation, allowed to
penetrate the specimen, and a 12-mm. number zero, round coverslip
was placed on the slide.
With the coverslip in place, generally heat from an alcohol lamp, would
expand the exoskeleton and float folded edges into position.
After 48 hours the coverslip could be ringed with asphalt.
Specimens which were used for examination of the fine structures and
trichobothria were mounted directly from 95% ethanol into CMC-10 and
allowed to "cure" for at least a week. This process, although slow, re-
tained the macrochaetae, trichobothria, and other fine structures in position.
It is highly desirable to use this technique because it permits preparation of
a large quantity of slides ina short time.
CHAETOTAXY
I. The Macrochaetal Pattern
The techniques referred to in the preceeding section permitted close
examination of the dorsal and lateral macrochaetae. The basic positions
of the dorsal macrochaetae and lasiotrichia described by Gisin (1964 and
1965) agree with the present observations. However, he did not include the
lateral setae, and placed the species he examined into species groups. I
*
Available from Turtox, General Biological Supply House, Inc.
Snider: The Chaetotaxy of Lepidocyrtus S. Str. 5)
have attempted to follow his examples wherever possible, but have departed
by showing a macrochaetal pattern for each species.
The constituents of the macrochaetal pattern include the macrochaetae,
lasiotrichia, and pseudopores. The macrochaetae (fig. 1) are pubescent and
arise from sockets surrounded by a chitinous ring. Lasiotrichia (Salmon,
1964) are long, fringed, sensory hairs (fig. 2) and are found on abdominal
segments II, I, andIV. Pseudopores (Gisin and DaGama, 1962) are struc-
tures which resemble pseudocelli, but lack internal structure (fig. 3). Gisin
and DaGama (1962) described a thin thread arising externally from the center
of the pseudopore. I have not as yet observed sucha structure. The pseudo-
pores occur on each body segment and Gisin (1963) reports that they also
occur on the coxae. Figure 4 is a composite chaetotaxal pattern and is repre-
sentative of all possible macrochaetae. A black dot indicates a macrochaeta,
an open circle (o) represents a pseudopore, anda short, curved line repre-
sents a lasiotrichium.
The species treated are arranged in order of their pattern complexity,
beginning with the most complex. Three basic regions of the trunk are
noted: the medial region (M), the paramedial (P), and the lateral region (L)
(fig. 4). These regions agree with the natural groupings of the macrochaetae
which are constant throughout the genus. All designated setae are indicated
by a segment number, region letter, and position subscript number. For
example, the first medial seta on abdominal segment IV is referred to as:
ABD IV, Mj). Lasiotrichia are designated by a "t" and a subscript number
such as: ABD IV, Mt9. By following this system, one is able to indicate
the presence or absence of a given seta by a notation. Setae below the
lateral edge of the body are not indicated, as they are beyond the scope of
the present study, as are also the setae of the parafurcular lobes and the
setae of abdominal segments V and VI.
Abdominal segment IV is the most variable in number and position of
setae. On the basis of macrochaetae, two distinct groups of species may be
discerned; group "A", with 3 medial macrochaetae; group "B", with two
medial macrochaetae. If the species are arranged under A and B groups
(table 1), they may be listed numerically from greater to lesser numbers
of macrochaetae.
TABLE 1.--Groups of species for North American Lepidocyrtus
based on the numbers of medial macrochaetae of ABD IV
"A" Group "B" Group
L. lignorum L. cinereus
~ unifasciatus ~ Januginosus
finensis heleni
curvicollis millsi
neofasciatus floridensis
paradoxus pallidus
violaceous cyaneus
Tables 2 and 3 summarize the macrochaetal pattern of the 14 species
examined. Thoracic segments II and III, and abdominal segment I have
only pseudopores, otherwise completely lack chaetotaxic elements, and
6 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
are therefore omitted from tabulations. It is significant to note that M, is
always present on ABD II and always lacking on ABD III; likewise, the para-
medials (P) are always lacking on both of these segments. Lateral seta (L,)
is consistently present on ABD II, but all other laterals are lacking on that
particular segment. All 14 species have the same pattern of lasiotrichia
which is as follows: ABD OU, Mt,, L,; ABD III, Mt,, Lt to; ABD IV, Pt,t
(fig. 4). Any unique positions or delétions are discussed under the individual
species.
TABLE 2.--Lateral macrochaetotaxal formulae of abdominal
segment III of Lepidocyrtus
Species
L. lignorum
gs
hs
unifasciatus
finensis
curvicollis
neofasciatus
paradoxus
violaceous
cinereus
lanuginosus
heleni
oe ee re et Ee a Beek ser
G6 OF 0) “29 09 00 GS Ute Go to 6
millsi
floridensis
po
oo
pallidus
Pee oe oboe eget oe Peak Bek
fh pk pp fuk kk kk fk pk ek peek
Din De eee ee ee ee ee
Oo Ow NO NO NNN NYO NY NY NY NY YN WD
cyaneus
Il. The Accessory Microchaetae Associated with
the Lateral Lasiotrichia of Abdomen II
Two lateral lasiotrichia and two macrochaetae (ABD III, t,t LL ) are
located on abdominal segment III. Associated with these structures are
microchaetae composing patterns which are species specific. I have adopted
a simple lettering system to designate the individual setae.
Lasiotrichium t, has a maximum of four pubescent microchaetae, commonly
arranged ina crescent shape, anterior in position to it. They are referred
to as A, a5 aga4. Commonly a, drops out of the pattern. Posterior to lasio-
ficken’ So additional microchaetae, b,b,, are found.
Similarly lasiotrichium t, has four microchaetae, CyCoC QC a” arranged in
a horizontal line drawn through the two lasiotrichia.
The Chaetotaxy of Lepidocyrtus S. Str.
Snider:
oT
ot
815
61, LT,91
61,,81,41,91
L191
8T 41,91
LT,91
815 9T
L191
615 LT,,91
8T AT 91
BT, 41,91
LT,,91
bs
ri
qo 4
ee ee ee ee eee eee eee ee,
rl
qo 4
ee ee ee ee ee ee ee
i
ee ee ee ee ee
ee ee ee ee
qd 4
VES ly
Peery
Bs ie og bap
As heda aa
beg hry
a he Ke
i
a ae ta
Prey Sq hy
Vb by by
Prey Ory bey
A he hs a
b
Sata
Te hela ha ls Ke aa
=
=
aa ae
Sa a SG
=
AA AY A
=
=
=
Py
OS > OS OOD "aD oe ey os Dd ey
De I cee A ce I cs I ce ce ce oe |
aoa Aa A AY A
snoueks
snptyred
‘STSUOPTIOTJ
“sy
“TueTey
snsouLsnuey
Snareurs
Snoaoe [OTA
snxopered
SNIVIOSeJOoU
ST[TOOTAINO
SISUOUT]
smieroseyiun
WINIOUSTT °
sotoedg
smaAdopidey jo AI JUSUISaS TeUTUIOPge JO IB[NUWIIO] TexeOLOeYIOIIVI[-- °F ATAVL
|
8 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
No microchaetae are known to be directly associated with macrochaeta
L,. However, two smooth microchaetae, one long (d,), and the other short
(5), are positioned mesad of macrochaeta L,.
Four possible microchaetae, lateral of the t,t,L,L, pattern, are referred
to as €,e,e,€,, in an anterior to posterior position. Occasionally there are
supplementary microchaetae associated with t, which are designated as s, and
S,. See figures 19-32 for positions and relative sizes of microchaetae. Table
4 indicates the comparison of the fourteen species examined.
III. The Accessory Microchaetae Associated with
the Anterior Lasiotrichia of Abdomen IV
Gisin (1964) described and named the microchaetae associated with the
anterior lasiotrichia of the fourth abdominal segment (ABD IV, P,t,) (figs.
33-36). He comments that the triangular pattern formed by these micro-
chaetae is not of particular significance. And the patterns for species over-
lap in many instances, allowing only grouping of species under the resultant
patterns. Gisin'’s system of notation is as follows: "One may distinguish one
external hair (e), one anterior (a) and one medial (m). In the center of the
triangle formed by the three hairs, one supplementary hair (s) is developed
," one (mp) medial posterior is present. This notation system for the
accessories has been followed.
CHAETOTAXY OF NORTH AMERICAN SPECIES
Lepidocyrtus lignorum (Fabricius), 1775
sensu Gisin, 1964
Figures 5, 19, 33
Lepidocyrtus lignorum (Fabricius) is easily confused with Lepidocyrtus
curvicollis Bourlet. North American taxonomists have designated, with few
exceptions, L. lignorum as L. curvicollis. Dr. Gisin (in litt.) informed me
that he thought L. lignorum was a common species compared to L.
curvicollis. After rechecking the determined collections of the Illinois Nat-
ural History Survey and Michigan State University, it was apparent that this
was indeed true. If color pattern alone is used to distinguish the species, it
is impossible to reliably separate the two. Lepidocyrtus lignorum is far
more common than L. curvicollis in North America.
CHAETOTAXY. --Lepidocyrtus lignorum has 29 setae on ABD IV (fig. 5),
the largest number of setae of the species examined. It is unique, being the
only species to have P, represented. The laterals are very similar to L.
unifasciatus from Le to L,, (table 3). S
The accessories of ABD Ill are simple plumose microsetae (fig. 19).
Microseta d, is longer thand,. This is a unique situation--the reverse is
true in all ntiaee species. Supplementary (s,) is very close to the first
lasiotrichia (t,). Table 4 indicates the microchaetal formula.
The accesSories of ABD IV consist of e, a, m, mp (fig. 33), the most
common pattern for the genus.
Snider: The Chaetotaxy of Lepidocyrtus S. Str.
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10 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
Lepidocyrtus unifasciatus James, 1933
Figures 6, 20, 33
This species is indigenous to North America. It can be distinguished
from other members of the genus on the basis of a deep blue band on abdominal
segment IV.
CHAETOTAXY. --L. unifasciatus has a macrochaetal pattern similar to
that of L. lignorum (fig. 6). A striking difference is on ABD III with the addi-
tion of L,, which is the only instance where that macrochaeta was observed.
Likewise, ei unifasciatus lacks ABD IV, PoP, which are found on L.
lignorum (table 3).
The accessories of ABD II (fig. 20) have the e series completely lacking,
whereas in L. lignorum e, and e, are present (table 4).
The accessories of Brice oF Mab IV (Ae. 33) are of the usual pattern, e, a, m, mp.
Lepidocyrtus finensis Maynard, 1951
Figures 7, 21, 33
Lepidocyrtus finensis was observed by Folsom in the 1920's, but never
described by him. Maynard collected the species in New York and published
his findings (1951). Records of L. finensis are very rare, suggesting that it
is an uncommon species. The specimens examined were collected in 1930 by
C, Aw Frost.
CHAETOTAXY.--L. finensis is close to L. — (table 3), according
to the macrochaetae. However, it has ABD IV, P, (fig. 7) which L. curvicollis
lacks.
The accessories of ABD III are similar to those in L. unifasciatus (table
4), having C. instead of c. Gigi2l).
The acobesorios of ABD IV are of the e, a, m, mp type (fig. 33).
Lepidocyrtus curvicollis Bourlet, 1839
sensu Gisin, 1964
Figures 8, 22, 35
As previously mentioned, L. curvicollis is considered to be close to L.
lignorum. Gisin (1964) could separate L. curvicollis once he discerned the
species, by the presence of pigment on thoracic segment II. The species is
not common in North America and previous determinations will have to be
checked before its range can be recorded accurately.
CHAETOTAXY. --The dissimilarity to L. lignorum, based on macro-
chaetae, is apparent at once (table 3). Lepidocyrtus curvicollis lacks
ABD IV P,P, and LeLiolis (fig. 8), but has LiLijg which is lacking on L.
lignorum.
The accessories.of ABD II have a simple pattern not found on L. lignorum
(table 4). Microsetae A5ae are absent, as well asd,d,. The supplementary
seta (s,) is present (fig. 294). The microsetae are strongly subclavate and
pubescent.
The accessories of ABD IV are unique in arrangement (fig. 35) and the
supplementary seta (s) is present.
Snider: The Chaetotaxy of Lepidocyrtus S. Str. Lt
Lepidocyrtus neofasciatus Wray, 1948, new status
Figures 9, 23, 33
When Wray (1948) described L. unifasciatus var. neofasciatus, he men-
tioned that there was some doubt concerning its systematic placement. James
who authored L. unifasciatus, checked the specimens and confirmed them as
L. unifasciatus var. Chaetotaxic studies have revealed that neofasciatus is
a valid species differing markedly from L. unifasciatus, and should stand
alone.
CHAETOTAXY.--L. neofasciatus lacks the macrochaeta ABD III L 4 found
on unifasciatus (table 3). It also has ABD IV P., which is not present on
unifasciatus, and the L series of ABD IV is mofe disjunct (fig. 9).
The accessories of ABD III lack dda, but have C5 and €, (table 4) which
are lacking on unifasciatus (fig. 23).
The accessories of ABD IV are of the e, a, m, mp type (fig. 33).
9
Lepidocyrtus paradoxus Uzel, 1890
Figures 10, 24, 34
L. paradoxus Uzel is one of North America's larger Collembola. Snider
and Fischer (1964) recorded its presence in North America for the first
time. European specimens for comparison were made available by Gisin,
and a rediscription from Michigan specimens was made. Goto (1953) de-
scribed L. christianseni as new, but Snider and Fischer's work has revealed
it to be a synonym of L. paradoxus.
CHAETOTAXY. --The macrochaetal pattern is similar to those of L.
neofasciatus, finensis and lignorum (table 3). The distribution of the L
series on ABD IV is disjunct (fig. 10).
The accessories of ABD III are similar to those of L. lignorum with ex-
ception of ABD IV dy, e,e, being absent on L. paradoxus. Both have sup-
plementary setae, s, for paradoxus and s, for lignorum (table 4). The
microchaetae are slightly clavate (fig. 24}.
The accessories of ABD IV (fig. 34) have the supplementary seta
(s) present.
Lepidocyrtus violaceous (Geoffroy), 1762
sensu Lubbock, 1873
Figures 11, 25, 33
There has been much discrepancy as to what constitutes the species
Lepidocyrtus violaceous (Geoffroy) and Lepidocyrtus cyaneus Tullberg. For
years taxonomists have confused the two species. Gisin (1964a) separated
them by using chaetotaxy and then substantiated his findings with more ob-
vious characters. The presence of scales on the first two segments of the
antennae and legs in violaceous permits its separation from cyaneus. This
character helped Gisin (1964b) to separate L. cyaneus var. assimilis Reuter
from cyaneus and to place it in synonomy under L. violaceous.
CHAETOTAXY.--ABD III, L, is missing on L. cyaneus, but is present
on L. violaceous (table 3). ABD, M, is present on L. violaceous, but
12 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
missing on L. cyaneus. The L series of ABD IV is the simplest of the A-
group (table 1), having 3 medial macrochaetae on that segment (fig. 11).
The accessories of ABD III (table 4) are the same as L. unifasciatus.
This is the only instance in the 14 species so far examined that repeats a
pattern (fig. 25).
The accessories of ABD IV are of the e, a, m, mp type pattern (fig. 33).
Lepidocyrtus cinereus Folsom, 1924, new status
Figures 12, 26, 33
Folsom (1924) found minor differences between his new variety, cinereus,
and L. cyaneus Tullberg s. str., but considered them insignificant. He com-
mented on the color difference: cinereus is metallic grey, while cyaneus is
deep blue. Notwithstanding he relegated cinereus to variety position. Gisin
(1964a) identified cyaneus, but did not mention the cinereus variety, and
later he was able to resurrect several valid species which had been placed
in synonomy with cyaneus. Based on the chaetotaxy and the differences
pointed out by Folsom, Lepidocyrtus cinereus Folsom is also a species dis-
tinct from cyaneus.
CHAETOTAXY. --The macrochaetae of ABD IV are the most complex of
the B-group (table 1). Lepidocyrtus cinereus also has ABD IV, Py (table 3)
which the others in the B-group are wanting (fig. 12).
The accessories of ABD III (table 4) lack the b and e series of micro-
chaetae (fig. 26).
The accessories of ABD IV are of the e, a, m, mp type pattern (fig. 33).
Lepidocyrtus lanuginosus (Gmelin), 1788
Figures 13, 27, 36
Lepidocyrtus lanuginosus (Gmelin) could be confused with lignorum
in size and color. Indeed, it is my belief that Maynard (1951) had the two
confused. Gisin (1964a) separated them on the presence or absence of scales
on the antennal segments J and II and legs. L. lanuginosus lacks those scales,
lignorum does not. y
This species is not common in the collections from North America ex-
amined by me.
CHAETOTAXY.--The macrochaetae pattern is somewhat similar to L.
floridensis n. sp. (table 3). Thoracic segment II has a lateral seta (L,).
ABD IV, P,P,P is an unusual combination in the B-group species (fig. 13).
The accessories of ABD III are reduced in number (table 4, fig. 27).
The accessories of ABD IV are of the e, a, m, mp type. However, e
is longer than normal, as in L. cyaneus (fig. 36).
Lepidocyrtus helenae, new species
Figures 14, 28, 33, 37-43
Body length up to 1 mm. The basic body color is violet, arranged on
the body segments and appendages as follows: antennal segment I dark violet,
segment II for the basal three quarters light violet merging on almost white
Snider: The Chaetotaxy of Lepidocyrtus S. Str. 13
at base, the apical quarter forms a dark ring, segment III violet, segment IV
violet; the precoxae and coxae violet, trochanter light violet, remainder of
leg white; furcula white; head dorsally with diffuse violet pigment becoming
darker in the genal area and very dark in the post-genal area, an anterior
black band connects the eye patches, ventrally the head is violet to light
violet; mesothorax up to and including abdominal segment II with dark violet
fascia, abdominal segment III sometimes with a small patch of lateral violet,
segment IV with a irregular violet fascia interrupted in the mid-dorsal re-
gion; parafurcular lobes dusted with small amounts of violet pigment (figs.
37 & 39); eyes 8 + 8 on dark patches (fig. 42); mouth parts prognathous with
a well developed molar plate on the mandible; antennal segments in the ratio
of 2: 6: 4: 8 (fig. 43); mesothorax enlarged, but not projecting into a cone;
ungues lanceolate with three inner teeth, unguiculus lanceolate with 4-5 small
teeth on its outer margin (fig. 41); a well-developed clavate tenant hair is
present; corpus of the tenaculum with one large irregular seta, rami with
four teeth (fig. 38); manubriun to dentes ratio 4: 5, dentes with many dorsal
crenulations; mucro typical of the genus with one apical tooth and anteapical
tooth with an anteapical spine pointed toward it from the basal position (fig.
40); scales on the head, trunk, and ventrally on the manubrium, lacking on
the legs from the trochanter to the claw and also not present on the antennae
--the usually fringed setae mixed with normal curving setae on the body.
CHAETOTAXY.--The macrochaetae are disjunct in the L series of ABD
IV (table 3, fig. 14).
The accessories of ABD III lack the b series and d series (table 4; fig.
28). | :
The accessories of ABD IV are of the e, a, m, mp type (fig. 33).
TYPES. --Holotype, Michigan, Monroe County, T75 R6E S824, June 17,
1965, collected by R. J. Snider. Paratypes, 8 individuals taken on the same
date as the holotype. Additional paratypes: Monroe Co., T75 R6E 824,
June 23, 1965, 2 specimens; June 29, 4 specimens; July 5, 2 specimens;
July 11, 1 specimen; July 17, 3 specimens; July 23, 38 specimens; August 4,
3 specimens; August 16, 5 specimens; August 22, 3 specimens; September 9,
2 specimens; all of which were collected by R. J. Snider in pit trap samples.
The holotype and paratypes are deposited in the Michigan State University
collection.
This species is commonly taken in small numbers with other members
of the genus in pit-trap samples. It is usually found in the litter of hardwood
forest floors where L. paradoxus, violaceous, unifasciatus and cyaneus occur.
I have collected L. helenae from June through September in Michigan. While
it does not constitute a large part of the collembolan population in the wood-
lots of southern Michigan, it is frequent in most samples collected.
It is my pleasure to name this species after Mrs. Helen M. Snider who
for sO many years put up with the "bug hunting" peculiarities of her son.
Lepidocyrtus millsi, new species
Figures 15, 29, 33, 44-50
Body length up to 1 mm. The basic body color is dark blue, arranged
on the segments and appendages as follows: antennal segment I light blue,
segment II pale basally with blue pigment on the distal quarter forming a
ring, segment III pale blue becoming dark blue on the distal quarter forming
14 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
a dark ring, segment IV pale blue becoming somewhat darker on the apical
half; precoxae and coxae blue, remainder of leg white; furcula uniformly
white; head dorsally dusted with light blue in some individuals, in others
white, an anterior dark band connects the eye patches and has an enlarged
medial spot, ventral side of the head may have some light blue dusting;
trunk segments from the second thoracic up to and including abdominal seg-
ment III always have dark blue fasciae, abdominal segment IV with a dark
blue fascia at the midpoint of the segment, the parafurcular lobes sometimes
dusted with light blue (figs. 44, 46); eyes 8 + 8 on dark patches (fig. 49);
mouthparts prognathous with well developed molar plate on the mandible;
antennal segments in the ratio of 2: 4: 3: 7 (fig. 50); mesothorax enlarged,
but not projecting anteriorly into a cone; ungues lanceolate with two inner
teeth, unguicula simple andlanceolate, a single weakly developed tenent hair
is present (fig. 48); corpus of the tenaculum with one large irreguiar seta,
rami with four teeth (fig. 45); manubrium to dentes ratio 4: 4.5, dentes with
many dorsal crenulations; mucro typical of the genus with one anteapical
tooth and apical tooth with a spine pointing toward the anteapical tooth from
its basal position; scales occurring on the greater part of the trunk, head,
and ventrally on the furcula, scales lacking on the antennal segments and on
the legs from the trochanter to the claw. The macrochaetae are fringed
and quite long, interspersed with short curving normal setae.
CHAETOTAXY.--The macrochaetae are simple in their pattern arrange-
ment (fig. 15). There is a gap between ABD IV, L, and L, . This is the
only species in the B-group to have the ABD IV, P,P P, eather (table 3).
The accessories of ABD III are more complex tn arrangement than the
other species of the B-group (table 4). Also, dd, are almost of the same
size (fig. 29).
The accessories of ABD IV fit into the e, a, m, mp type pattern (fig. 33).
TYPES. --Holotype, Putnam County, Springdale, March 19, 1927, col-
lected by T. H. Hubbell. Paratypes, 20 individuals taken on the same date
as the holotype. The holotype and paratypes are deposited in the [llinois
Natural History Survey collection. Five paratype specimens are in the
Michigan State University collection.
This species could easily be confused with Lepidocyrtus helenae but
may be separated by the nature of the claws. lL. helenae has three inner
teeth on the unguis, while millsi has two. Also, helenae has a serrated
outer margin on the unguiculus which makes it unique.
I take pleasure in naming this species after Dr. Harlow B. Mills, who
has been so helpful during the beginning years of my efforts in Collembola
taxonomy.
Lepidocyrtus floridensis, new species
Figures 16, 30, 34, 51-57
Length of body up to 1 mm. The basic body color is white with dark
blue-black maculations laid down in flecks of pigment distributed as follows:
antennal segment I light blue throughout, segments II and ITI light blue be-
coming darker apically, segment IV uniformally light blue; legs entirely
white except coxae frequently with a light dusting of dark blue which con-
tinues over basal fourth of the trochanter; furcula white; ventral side of the
head with a light dusting of dark-blue black pigment, the dorsal side with a
Snider: The Chaetotaxy of Lepidocyrtus S. Str. 15
dark anterior band connecting the eye patches; mesotergum and metatergum
with a dark blue-black lateral macula; the first segment of abdomen with
blue-black fascia interrupted mid-dorsally with white, sometimes continuous;
abdominal segment IV with a blue-black fascia, frequently discontinuous to the
extent that only a patch may be seen laterally on either side of the segment
(figs. 51, 52); eyes 8 and 8 on dark patches (fig. 55); mouthparts prognathous,
with a well-developed molar plate on the mandible; antennal segments in the
size ratio of 2: 3: 3: 6 (fig. 54); mesothorax enlarged, but not projecting
anteriorly into a cone; ungues lanceolate with an outer basal tooth and three
inner teeth, unguicula with four outer teeth and lanceolate, a single clavate
tenent hair (fig. 56); corpus of the tenaculum with one large irregular seta,
rami with four teeth (fig. 53); manubrium and dentes in the ratio of 1: 1, with
many dorsal crenulations on the dentes; mucro with large apical tooth anda
basal spine pointing toward anteapical tooth (fig. 57); scales on the greater
part of the body and on the ventral surface of the furcula, scales not occurring
on the legs from the trochanter to the claw, or on the first two segments of
the antennae. The basic setae of the body are fringed and are heavy on the
legs.
CHAETOTAXY. --The macrochaetae of ABD IV lacks P, which is found
on all other species described in this paper (table 3). The © series show a
disjunct pattern (fig. 16).
The accessories of ABD III (table 4) are very numerous, only the sup-
plementaries are missing. The microchaetae are subclavate (fig. 30). The
seta d, is missing.
The accessories of ABD IV resemble L. paradoxus (fig. 34).
TYPES. --Holotype, Florida, Stock Island, December 29, 1951, col-
lected by R. Richards and L. Stannard. Paratypes, 108 individuals col-
lected on the same date as the holotype. Additional paratypes were col-
lected from the following Florida localities: Sanibel Island, April 26, 1927,
M. D. Leonard, collector, 16 specimens; Everglades National Park, Royal
Palm Ranger Station, December 27, 1951, Richards and Stannard, collectors,
21 specimens; Key West, December 27, 1951, Richards and Standard, col-
lectors, 5 specimens. The habitat for this species is grass clumps and
ground litter.
Holotype and paratypes are deposited in the Illinois Natural History Sur-
vey collection. 20 paratypes are also in the Michigan State University col-
lection.
Lepidocyrtus pallidus (Reuter), 1890
sensu Gisin, 1965
Figures 17, 31
Lepidocyrtus pallidus (Reuter), originally considered a valid species,
has commonly been placed into the cyaneus complex as a variety. Gisin
(1965), using chaetotaxy and morphological characters, resurrected the
species. His dorsal chaetotaxy drawing of the body associate it with L.
lanuginosus and cyaneus. The species has been further defined in this
study.
CHAETOTAXY. --The macrochaetal pattern is similar to L. cyaneus,
but differs on ABD IV lacking L,L,. and by having in addition L, 41.
(table 3; fig..17).
16 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
The accessories of ABD III lack the d and e series (table 4) and supple-
mentaries (fig. 31).
The accessories of ABD IV are of the e, a, m, mp type pattern (fig. 33).
Lepidocyrtus cyaneus Tullberg, 1871
Figures 18, 32, 36
Having been confused with L. violaceous for so many years, L. cyaneus
was often misidentified in North America until Gisin (1964) redefined cyaneus.
I believe that cyaneus is not as common as violaceous in our fauna. The
presence of scales on antennal segments I, II and the legs separate cyaneus
from violaceous.
CHAETOTAXY. --Thoracic segment II bears a lateral macroseta (L,)
(fig. 18). ABD III L, is missing. The macrochaetae of ABD IV are the
least numerous of alP 14 species considered here (tables 2 and 3).
The accessories of ABD III lack the c, d, and e series as well as supple-
mentaries (table 4). The microchaetae are subclavate and small (fig. 32).
The external accessory (e) of ABD IV is almost twice the normal length
(fig. 36), similar to L. lanuginosus.
SUMMARY
The mounting technique described in this work proved useful in making
microscopic examinations of 14 species of Lepidocyrtus s. str. found in
North America. whoa
The macrochaetal patterns of the species were studied and illustrated.
Two major groups of species were discerned within the genus on the basis
of macrochaetae. Microchaetae (accessories) were described from abdominal
segments III and IV. The combination of macro- and microchaetal patterns
characterize the species studied and give foundation to the definition of in-
dividual species. Chaetotaxy is not considered as an end in itself, but rather,
an aid for the identification of difficult species.
Chaetotaxy was used in identification of three new species; Lepidocyrtus
helenae, Lepidocyrtus millsi, and Lepidocyrtus floridensis. Specific status
for two varieties was proposed; Lepidocyrtus neofasciatus Wray and
Lepidocyrtus cinereus Folsom.
It is hoped that future work of a revisionary nature on the group will be
made easier and be given a more uniform method of presenting chaetotaxic
data as a result of this study.
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Massoud, Z. 1964. Description d'un nouveau genre de poduromorphe
(Collembola: Arthropleona). Rev. Biol. Ecol. Sol., 1: 511-518.
Maynard, E. A. 1951. A Monograph of the Collembola or springtail
insects of New York State. Comstock Publ. Co. Inc., Ithaca. 339 p.
Mills, H. B. 1933. A monograph of the Collembola of Iowa. Collegiate
Press, Inc., Ames. 143 p.
Murphy, D. H. 1966. Taxonomy and bionomics of eight species of
Sphaeridia (Collembola, Sminthuridae) in a Singapore flood-plain.
Rev. Ecol. Biol. Sol., 3: 65-95.
Salmon, J. T. 1964. An Index to the Collembola. Roy. Soc. New Zealand,
pts. 1-2, (Bull. 7) 1-644.
18 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
Snider, R. and R. Fischer. 1964. A Palearctic springtail, Lepidocyrtus
paradoxus Uzel, found in North America (Collembola: Mydontidae)
Trans. Amer. Micr. Soc., 83: 86-89.
Wray, D. L. 1948. Some new species and varieties of Collembola from
North Carolina. Bull. Brooklyn Ent. Soc., 32: pp. 44-53.
Yosii, R. 1956. Monographic zur Hohlencollembolen Japans. Contr.
Biol. Lab. Kyoto Univer., 3: 1-109.
1959. Collembolan fauna of the Cape Province, with special
reference to the genus Seira Lubbock. Biological Results of the
Japanese Antarctic Research Expedition 6. Special Pbl. Sets. Mar.
Biol. “have: 23 pi
1960. Studies on the Collembolan Genus Hypogastrura. Amer.
Mid. Nat., 64: 257-281.
1961. Phylogenetische bedeutung der Chaetotaxie bei den
Collembolen. Contrib. Biol. Lab. Kyoto Univ., No. 12, 37 p.
19
Snider: The Chaetotaxy of Lepidocyrtus S. Str.
©
ie.
)
=
©
>
©
Q
Fig. 1.--Macrochaeta. Fig. 2.--Lasiotrichium. Fig. 3.--Pseudopore.
Fig. 4.--Dorsal composite view of Lepidocyrtus indicating the regions of
the body and positions of the macrochaetal elements.
Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
20
“aeyevyooroeul Testop “‘o‘u ‘Avi M Snyetosejoou snyzAoopidey--°¢ “Sly ‘“aejevYDOIOeUI TeSAOp “aTINOg ST][OOTAIND
snjtAoopidey--°g “Sly ‘aevjaeyooroeu [esiop ‘preuAdey stsuouly smprAd0pidey--*), ‘31q ‘aejaeyooroeU Tes.op
‘Somer snyetosejiun snyzAoopidey-- "9 “sIy ‘aejaeyooroeU [essop ‘(Ssntoliqe,) wWnIOUsTT SmaADOpIdey--"¢ *31q
6
:
8
L 9 G
ee
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O
“avjavyooIOe Ww
Tesiop ‘*ds *u sevuatay snyzAd0ptde7q-- "PL ‘SIT ‘avjovyooroeU Tes.0p ‘(UITaWIy) SnsouTSnue, smaAd0pIde'7
--"§]T “SIT ‘avjaeyooroeu [esiop ‘*o-u ‘wIOSsToO,J Sne1euto SnytAdopidey-- "Zl “sly ‘eevjaeyooroeUl Teszop
‘(Aorjjoay) snosovjota sm.aAd0ptdey-- ‘IT “SIT ‘avjaeyoor1oeu [esi1op ‘Tazq snxopeied snyztAd0pidey-- oy “317
21
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Snider: The Chaetotaxy of Lepidocyrtus S. Str.
Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
22
‘avJOVYOOINVUI [eSIop ‘sraeq iin, sneuvAod smaAhooptdeT
--"Q] ‘SILT ‘ovjaeyoor1oeUl [Tesrop ‘(19jyney) Ssnpti[ed smaAdopidey-- "pT “3lq ‘aejyaeyooLoeW Tes.iop
‘‘ds ‘u stsuapliopy snjyzA00pidey--‘9] “SIq ‘aevjyovyooroeu Tesaop ‘‘ds ‘u tsyqtum smazAoopidey-- ‘cy *31q
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Snider: The Chaetotaxy of Lepidocyrtus S. Str. 23
a
Als
is
A
19. rl ©
4.
24.
The Microchaetae (Accessories) of ABD III. Fig. 19.--L. lignorum. Fig.
20.--L. unifasciatus. Fig. 21.--L. finensis. Fig. 22.--L. curvicollis.
Fig. 23.--L. neofasciatus. Fig. 24.--L. paradoxus.
Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
| ©
25, | 28.
va ©
Pe a |) N
9 Oo yeh 30 \
The Microchaetae (Accessories) of ABD III. Fig. 25.--L. violaceous. Fig.
26.--L. cinereus. Fig. 27.--L. lanuginosus. Fig. 28.--L. helenae. Fig.
29.--L. millsi, Fig. 00 ++h. cioridensis.
Snider: The Chaetotaxy of Lepidocyrtus S. Str.
E \ )
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© \ ©
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34. 36.
The Microchaetae (Accessories) of ABD III. Fig. 31.--L. pallidus. Fig.
32.--L. cyaneus. The Microchaetae (Accessories) of ABDIV. Fig. 33. --
es ——— —_
re
helenae, millsi. Fig. 34.--L. paradoxus, floridensis. Fig. 35.--L.
curvicollis. Fig. 36.--L. lanuginosus, cyaneus.
29
26 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
Snider
Lepidocyrtus helenae n. sp. Fig. 37.--L. helenae, dorsal view habit
drawing. Fig. 38.--Tenaculum with seta. Fig. 39.--L. helenae, lateral
view habit drawing. Fig. 40.--Mucro. Fig. 41.--Right claw of third leg.
Fig. 42.--Eye patch of left side of head. Fig. 43.--Antenna.
Snider: The Chaetotaxy of Lepidocyrtus S. Str. 27
ou,
49.
48.
Snider
Lepidocyrtus millsin. sp. Fig. 44.--L. millsi, dorsal view habit cohae:
(drawing by J. W. Folsom). Fig. 45. =Tenaculum with seta. Fig. 46. -
L. millsi, lateral view habit drawing. Fig. 47.--Mucro (after Folsom).
Fig. 48. -- =-Right claw of third leg (after Folsom). Fig. 49.--Eye patch of
left side of head (after Folsom). Fig. 50.--Antenna.
28 Contr. Amer. Ent. Inst., vol. 2, no. 3, 1967
Sader
Lepidocyrtus floridensis n. sp. Fig. 41.--L. floridensis, lateral view habit
drawing. Fig. 52.--L. floridensis, dorsal view habit drawing. Fig. 53. --
Tenaculum with seta. Fig. 54.--Antenna. Fig. 55.--Eye patch of left
side of head. Fig. 56.--Right claw of third leg. Fig. 57.--Mucro.
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Contributions
of the
American Entomological Institute
Volume 2, Number 4, 1968
~_
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA. - IT.
he genus Aedes, subgenus Neomacleaya Theobald in Southeast Asia.
by
Mercedes D. Delfinado
ee +4 ey iA
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Contributions
of the
American Entomological Institute
Volume 2, Number 4, 1968
~<
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA. - If
genus Aedes, subgenus Neomacleaya Theobald in Southeast Asia.
by
Mercedes D. Delfinado
CONTENTS
ENP ROIUC TION ce a ee ee i ee ee ee we 1
SPECIES OF THE SUBGENUS NEOMACLEAYA KNOWN FROM
SOULE BOI i oe ra a a ee ee 2
KEYS TO THE SPECIES IN SOUTHEAST ASIA
I ae a ae ee 3
A a re ee ee oe ae 7
ARV AG Cia ee hee ks Re ae ee ee 11
DESCRIPTIONS OF THE SPECIES
CO OVIOS VIUS TATOO 6 ek ee ee ee ee 13
Fre Ee) a a eae 15 ®
Waves Vins DBHOOn ee a ee bees 16 4
Pe resG OMA bk a ek ae eo ee 18 @
GOS TU ee ee ee ee ak eae OE 19
FORCES Te BOWE fo Ne Ce ew le Se eee 21
DOYS era ee ee ee a a a ae ag 22
macroaixou Dyar and Shannon... 6 6 606 6 ee ke ks 23
moavedy sen Dyer and Shannon... 6 kk ek kk eh ee we 25
neomacrodixoa King and Hoogstraal. ......... . .40
Nieves POW) a ea a ae 28
WOO EO a a ee eS ae oe 30
Wo a ek a ee 31
POOH s TAIOW Se Ca a es Ss a eS Oa ee 32
PRT ODE AS7 se UO SOR ag ee a Se ee 33
Oe ets OL a ce ae eS ae ce ee 34
VOCUS HBO. Po ea ee Rk oe ee ee ee ee 35
PODEFIED TOON ee ee See ee 8 ee ee Oe 35 8
SQ oensis AOSD. SU ae a ee ee 36
ep avie (heieertery ee ea ae 38
Ceres ice ee a a 39
Cee Tiler i a oe ee ee ee 41
SPECIES OUTER NCA ee ee a ee Oe ee
CORRECTIONS TO AEDES (NEOMA CLEAYA) IN THAILAND. ..... 42
ACKNOWLEDGE WENT 6256 ee ia ie we eee eee 42
LIVE et ee CE a i le el ee ee ee eee 42
APPENDIX - PRESENT STATUS OF THE AEDES (NEOMACLEAYA)
FAUNA OF BOU FRE AST ABTA 6 cee ee we ee ee 44
BN Re aise) el uaeboew 8 ae cele 47
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. II.
THE GENUS AEDES, SUBGENUS NEOMACLEAYA THEOBALD
IN SOUTHEAST ASIA. *
By
Mercedes D. Daiticada
INTRODUCTION
This is the second of two papers dealing with a revision of the Aedes
subgenus Neomacleaya of Southeast Asia. The previous paper (Delfinado
1967) described the species represented in Thailand, the present one treats
all of the remaining species known to occur in Southeast Asia and an addi-
tional new species from Thailand. The name Neomacleaya Theobald was
resurrected from synonymy with subgenera Aedes s. str. and Verrallina
by Delfinado (1967). It was recognized as a distinct subgenus for the primar-
ily Oriental species formerly placed in Aedes or Verrallina because it has
little in common with the type species of subgenus Aedes (cinereus Meigen,
described from Europe) but resembles butlevi Theobald, the type species of
Verrallina. The Neomacleaya are distinguished from Vervallina by the re-
markable development of the female and male terminalia, and by certain
characters of the immature stages. The subgenus is restricted in distribu-
tion to the Oriental region except A. panayensis and A. neomacrodixoa,
which also occur in the Australasian region.
The material used in this study is from the collections of the U. S.
National Museum, Washington, D. C. (USNM); B. P. Bishop Museum,
- Honolulu (Bishop); British Museum (Natural History), London (BM); California
Academy of Sciences, San Francisco (CAS); Academy of Natural Sciences,
Philadelphia (ANS); Chicago Natural History Museum (CNHM); Johns Hopkins
School of Tropical Medicine and Hygiene, Baltimore (JHS), and the Instituut
voor Tropische Hygiene, Amsterdam (AM). The type specimens of the new
species will be deposited in the respective institutions as indicated in the
type data.
. This work was supported by Research Contract No. DA-49-193-MD-2672
from the U. S. Army Medical Research and Development Command, Office
of the Surgeon General.
2 South East Asia Mosquito Project, Smithsonian Institution, Washington,
D. C. 20560.
2 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
Of the 55 species in Neomacleaya, 42 are presently known in South-
east Asia. It seems that in this subgenus there are several widely distribute
dominant species and a number of specialized endemics. It is probable that
there are still several unrecognized species within its geographical range.
Very little information on adult bionomics is available though some species
are known to take blood; little is also known of the specific larval habitats,
- and in many species one or more stages are unknown. Much collecting and
individual rearings of several species remain to be done.
Four new species from the Philippines, Malaya, North Borneo and
Thailand respectively are added here. Aedes malayi (Leicester) is here
placed as a synonym of A. fragilis (Leicester) and Aioretomyia perdita
Leicester is treated as a species inquirenda.
The terminology, larval and pupal chaetotaxy and format used are
as given in the previous paper. Nothing is added to the subgeneric account
which I have already discussed (Delfinado 1967): :
During the present study the diagnostic value of the structure of the
tarsal claws was revealed and hence this aspect receives more than usual
prominence here.
The pupae as presently known show few reliable specific characters
hence a key for their identification is not given.
SPECIES OF THE SUBGENUS NEOMACLEAYA KNOWN
FROM SOUTHEAST ASIA.
In this list species marked with a single asterisk (*) are dealt with
here in detail, those which are unmarked have received formal treatment in
Delfinado (1967). A double asterisk (**) following a country indicates a new
record.
1. andamanensis Edwards - Andaman Islands, Philippines, Thailand,
India, Malaya, Sumatra, Java, North
Borneo, Vietnam
2. atrius Barraud - India (Assam), Thailand
3. campylostylus Laffoon* - Philippines
4. cautus Barraud - India, Thailand, North Borneo, Malaya
5. clavatus Barraud - India, Thailand, Vietnam
6. cretatus Delfinado - Thailand
7. cyrtolabis Edwards - Singapore, Thailand
8. dermajoensis Brug - Sumatra, Thailand
9. dux Dyar andShannon - Philippines, Andaman Islands, Thailand,
Java, Hainan Island, Vietnam
10. fragilis (Leicester)* - Malaya
11. gibbosus Delfinado - Thailand, Java, Malaya
12. hamistylus Laffoon* - Philippines, North Borneo**
13. hispidus Delfinado - Thailand
2(1).
Delfinado: Aedes (Neomacleaya) in Southeast Asia 3
incertus Edwards - Thailand, Java, Malaya, North Borneo
indecorabilis (Leicester) - Malaya, Thailand, North Borneo
indicus (Theobald) - India, Thailand, Ceylon
ishigakiensis Bohart* - Ryukyu Islands
johnsoni Laffoon* - Philippines
latibennis Delfinado - Thailand
leicesteri Edwards* - Malaya, North Borneo**
lugubvyis Barraud* - Malaya**, Burma, Andaman Islands
macrodixoa Dyar and Shannon* - Philippines
margarsen Dyar and Shannon* - Philippines
neomacrodixoa King and Hoogstraal* - Celebes, New Guinea
nigrotarsis (Ludlow)* - Philippines
notabilis Delfinado - Thailand
nubicola Laffoon* - Philippines
pahangi n. sp.* - Malaya
panayensis Ludlow* - Philippines, New Guinea, Schouten Island,
Morotai, Moluccas
philippinensis n. sp.* - Philippines
prioekanensis Brug* - Sumatra
protuberans Delfinado - Thailand
pseudodiurnus (Theobald) - India, Thailand
varus n. sp.* - North Borneo
robertsi Laffoon* - Philippines
siamensis n. sp.* - Thailand
singularis (Leicester)* - Malaya, Sarawak
torosus Delfinado - Thailand
uncus (Theobald) - Philippines, Thailand, Java, North Borneo,
Malaya, Sarawak, India (Assam)
vallistris Barraud - India (Assam), Burma, Thailand, Cambodia**
varietas (Leicester)* - Sumatra, Java, ** Malaya
virilis (Leicester)* - Malaya
KEYS TO THE SPECIES IN i 4 dl eae ope ASIA
FEMALES
Hind.tarsal claws toothed (Figure 26)... 2 04.5. 004 Ones i 3G 2
Hind tarsal elaws simple (Figure 17)..j%. < scsi... 8
Abdominal terga dark, without pale markings. ........... 3
Abdominal terga with pale markings (Figures 1, 2, 4, 5). ..... 6
1 Females of the following unknown: cyvtolabis, dermajoensis, indecorabilis,
leicesteri, nubicola, prioekanensis, protubevans, pseudodiurnus, varus, torosus
and
virilis,
4(3).
5(4).
6(2).
7(6).
8(1).
9(8).
10(9)..
11(10).
Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
Postatrial plate with a pair of hairy lobed structures; preatrial
plate consisting of 2 small, hairy separate parts; spermathecae
each with a emall neck Ve a notabilis
Postatrial plate not as above, without hairs; preatrial plate entire;
spermathecae without distinct necks.............0.¢ A
Spermathecae reticulate; terminalia as in figure 23. .........
Spermathecae smo, Oe ee ee Pe ee 4)
Postatrial plate ovate with a rather large opening (Figure 25)... .
: robertsi (p. 35)
Postatrial plate quadrate with a small opening (Figure 9).......
hamistylus (p. 16)
Vertex with a patch of white scales in front extending downwards to
frons; terminalia as in figure 26. ...... siamensis (p. 36)
WEttes NOU AS SDOVE. 6 OR he Ce ee PR Sa a 7
Terminalia with a markedly large preatrial plate expanded laterally
(Figure 12) cee ee oe Leer, ishigakiensis (p. 18)
Terminalia with a rather small preatrial plate without lateral
expansions (Ficure 8). eV eS ee a fragilis (p.15)
Abdominal terga dark brown or with light brown lateral patches,
sometimes a small lateral spot of almost whitish scales is
present on tergum I (Figure 3); terminalia fairly simple, post-
atrial plate with a pair of hairy finger-like processes. .....
aux
Abdominal terga with distinct pale markings; terminalia not as
PENT Oe a ee a ee ere oe a 9
At least one of the terga with a complete pale band. ........ 10
Terga with pale lateral patches, sometimes produced on to dorsum,
rarely complete in margarsen (Figures 1, 4, 5)........ 14
Only tergum II with a complete pale band; terminalia with postatrial
and preatrial plates fairly simple (Figure 13). ..........
lugubris (p. 22)
Terga II-VII with complete pale bands (Figure 2); terminalia not as
ROOK S a ee 11
All spermathecae with narrow necks. ........6eseee08-8 12
Either 1 or 3 spermathecae with swollen necks.......... 13
ae
12(11).
13(11).
14(9).
15(14).
16(15).
17(16).
18(17).
19(17).
20(14).
Delfinado: Aedes (Neomacleaya) in Southeast Asia 3)
Postatrial plate with broad wing-like structures laterally; preatrial
plate consisting of widely separated, long, narrow parts... .
cretatus
Postatrial plate not as above; preatrial plate narrowly separated,
consisting of very large, ovate parts. .... hispidus
Only the largest spermatheca with a bent, swollen neck; preatrial
plate partially sclerotized. .......... indicus
Three spermathecae each with a large, bulbous neck; preatrial
plate heavily sclerotized (Figure 21). . . . nigrotarsis (p. 28)
Lower mesepimeron with fine hairs (Figures 2, 4, 5). ..... 15
Lower mesepimeron bare (Figures 1, 3)............6. 20
Sternopleuron with about 20 strong bristles behind pale scale patch;
spermathecae without necks;terminalia as in figure 22. .....
singularis (p. 38)
Sternopleuron with 10 or less bristles. .............6. 16
Metameron with fine hairs; postatrial sclerite convex medianly
with strongly lobed lateral corners; postatrial plate with scal-
loped distal margin of opening. ....... gibbosus
Metameron Dare. 66 i ee ws Re es 17
At least one spermatheca with a large, swollen neck. ...... 18
All spermathecae with small necks. ............0.008-4 19
Usually only the largest spermatheca with a large, swollen neck;
terminalia as infigure15....... neomacrodixoa (p. 2'7)
Three spermathecae each with a swollen neck; terminalia as in
UTS T4 g e a a we e e macrodixoa (p. 23)
Postatrial plate simple without hairs; preatrial plate consisting of
2 small poorly sclerotized parts weakly connected medianly
(Pigure 29 iid ke oe Plo eas panayensis (p. 32)
Postatrial plate with a large cordate opening, many long hairs;
preatrial plate large, weakly divided, narrow basally. .....
UNCUS
Metameron with fine hairs (Figure 1); postatrial plate with bluntly
pointed distal arms (Figures 17, 19) (see figures for structural
CINCTCRER Re a a Ba andamanensis
margarsen (p. 25)
johnsoni (p. 20)
MICTAIMOPON DRT Ce Or eee Oe Oe PIG 21
2.1(20).
22(21).
23(22).
24(23).
25(23).
26(25).
27(25).
28(27).
Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
Postatrial plate pocket-shaped, without hairs; preatrial plate
poorly sclerotized, consisting of 2 small lobes narrowly
Conmmected distally. 6 ci ek ee ee eae incertus
NOU GS ADOVED Sh. ko er a ee ee ee 22
Terminalia with preatrial plate consisting of 2 widely separated,
Iong, narrow, parallel-sided parts (Figure 24).........
philippinensis (p. 33)
INOU GS QDOVE. 3. oe ee a re Poe 23
Postatrial sclerite with lobed lateral corners. ......... 24
Postatrial sclerite rounded... 1.0. toe ew 25
Preatrial plate large, subquadrate, weakly divided, hairy (Figure
BOOS Be a a varietas (p. 40)
Preatrial plate consisting of 2 narrow, long parts with hairy
rounded ends and connected at this portion. . . cautus
Postatrial plate with distal arms. . . . 6.4 ee ee ee 26
Poptatrial plate without arms... ON at
Distal arms short and straight; preatrial plate consisting of 2
small separate parts. .....-....2+.-. vallistris
Distal arms incurved, fairly broad; preatrial plate consisting of
2 large separate parts (Figure 7). . . . campylostylus (p. 18)
Postatrial plate lightly sclerotized, covered mostly with short,
Branched haivs 2 easy ee as latipennis
Postatrial plate with many coarse, long hairs only around margin
OP OpCNIMNE eee Ga ke 28
Preatrial plate weakly sclerotized at middle, with a pair of
sclerotized transverse ridges and a pair of additional cres-
centic platelets on each side. ......... clavatus
Preatrial plate consisting of 2 large, subquadrate parts narrowly
contiected distally so.) ol. er atrius
Delfinado: Aedes (Neomacleaya) in Southeast Asia t
MALES!
1. Abdominal terga wholly dark, or with pale brown lateral patches
or a small lateral spot of almost whitish scales may be
present on tergum I (Figure 3): 6 2 2 6 ee ek 2
Abdominal terga with pale markings. ..............6. 8
2(1); .. Hind tarsal claws simple (Figures 13, 15, 28, 24) ........ 3
Hind tarsal claws toothed (Figures 8, 10, 11, 18, 26)....... 4
3(2). Basimere projecting laterally towards inner margin, with a sub-
apical process and a short curved spine. ..... dux
Basimere without subapical or apical projections but with a patch
of strong spines and setae on inner margin (Figure 24). ....
philippinensis(p. 38)
4(2). | Basimeres separated, with a short blunt apical projection and 2
large spines on inner margin near base (Figure 8). .......
prioekanensis (p. 34)
Basimeres joined subapically: . se 4G eer ee
5(4). Vertex with a patch of white scales in front extending downwards to
frons; terminalia as in figure 26...... siamensis(p. 377)
Vertex 110. 4s above.) 6 icy ee Pe a a eS 6
6(5). Fore tarsal claws unequal, toothed, the smaller minutely so; mid
claws unequal; only the smaller claw toothed (Figure 10);
terminalia as in figure 10. ......... hamistylus (p. 17)
Fore and mid claws unequal, only the larger claw toothed (Figure
13) hs eee ee ee ee a 7
7(6). Terminalia as in figure 13, basimere rounded apically bearing a
small, blunt, curved process. ........ robertsi (p. 36)
Basimere bluntly produced distally, without such process; disti-
mere markedly swollen at middle, abruptly narrowed and
curved distally, with many setae at the swollen portion. ....
indecorabilis
8(2). Fore and mid claws equal or unequal, each claw toothed (Figure
1 vfales of the following unknown: /ispidus, notabilis, pahangi; incertus and
indicus uncertain.
10(9).
11(10).
12(11).
13(8).
14(13).
15(13).
16(15).
17(16).
18(16).
Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
Hind claws toothed; basimeres joined subapically, with an apical
projection bearing a large, hooked spine (Figure 11)......
ishigakiensis (p. 18)
Hind claws simple; basimeres separated. ...........e.e. 10
Fore and mid tarsal claws equal; basimere with a narrow, flattened
apical projection with serrated tip (Figure 23). .........
panayensis (p. 32)
Fore aud miid claws anegual ee OE ION A a 11
Basimere without apical projection but with 2 large spines near
base and a median flattened projection with serrated tip; para-
proct very small (Fieure 24)/o.00 ee 24 varus (p. 35)
Basimere with short differentiated apical projections; with long,
Weer Para OC. is a ke Se ec ew eae on ae 12
Basimere projected apically into a short pointed process with 1-2
subapical spines and 2 long basal spines. ............
dermajoensts
Basimere with 2 differentiated apical projections, one of which is
long and slender, the other short and hairy, a large peg-like
process and 1-2 large spines near base. . . .cretatus
At least one of the terga with a complete pale band. ........ 14
Terga with pale lateral patches only sometimes produced on to
GCGVBUIS Se a Ne EP Pe ye 15
Only tergum II with a complete pale band; terminalia as in figure
TO SI ee ae OO eS lugubris (p. 22)
Terga II-VUI with complete pale bands; terminalia as in figure
a ea ee nigrotarsis(p. 28)
Lower mesepimeron with fine hairs. ..........4.++e62.6-. 16
Lower mecepimeron bare. . fe a A we we 25
Tergum IX with a markedly large median Y-shaped structure. . .17
PGR ee ee a ee ee ei es 18
Each hairy branch of the Y-shaped structure expanded and with a
lateral lobe (Figure 15). ........ neomacrodixoa(p. 27)
These branches wider and more abruptly tapered distally and with-
out a lateral lobe (Figure 15). ...... macrodixoa (p. 24)
Parenroct 100m ONC Paper ec. bo eke a a ee a 19
Paragproct very SNOriis, bee PI Ne 22
19(18).
20(19).
21(20).
22(18).
23(22).
24(23).
25(15).
26(25).
Delfinado: Aedes (Neomacleaya) in Southeast Asia 9
Tergum IX projecting medianly into a wedge-shaped structure;
paraproct unusually long and wavy. .... cyrtolabis
Not a8 Sboves oo Sic ee a a a re 20
Distimere bifurcate at distal half; sternum IX projecting distally,
with a deep median concavity (Figure 22). . singularis(p. 39)
Not 868 ANOVO. ee ee ee a ewe ee a eS 21
Distimere short and broad for most of its length and with a pointed
lateral projection at apex; apical projection of basimere with
5 large spines and a small inner process (Figure 22)......
nubicola(p. 30)
Distimere very long, tapered distally, with a small pointed
process at the middle; apical projection of basimere without
spines (Figuve 22)c) erie oa Oe ee ae virilis (p. 41)
Sternum IX with 3 or 4 large marginal spines; basimere with 2
long slender projections, one of which bears 2 or 3 spines at
apex while the other is split at tip, also with a short process
armed with 6-8 strong spines and a shorter blunt projection.
uncus
Not aS abovessii i eas EAU ae as a a 23
Distimere much enlarged and broadened distally; basimere with a
large, curved spine and a short subapical projection armed
with 2 large and 3 small spines (Figure 8)............
leicesteri(p. 21)
Distimere swollen at the middle, tapered distally. ........ 24
Basimere with a long, pointed apical projection and 2 short pro-
cesses, one of which bears 3 or 4 sharp spines, the other
simple and potted) gis cee Ww protuberans
Basimere with a large, hairy lobe, a long slender projection and
large hooked spine at inner margin. ..... tovosus
Metameron. with tine hares. yt A151 < Oe Sa ow 26
MetAIMeCrOn DETER eee. ws 29
Basimere projecting distally then abruptly narrowed and bent before
tip, with a short process on inner margin bearing a large spine,
a subapical lobe with 3 or 4 spines; distimere stout and ex-
panded Gistallyei ys wl we wlik a ON Ws gibbosus
Basimere with forked apical projections; distimere slender. . . 27
10
27(26).
28(27).
29(25).
30(29).
31(30).
32(31).
33(30).
34(33).
Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
Basimere with the apicodorsal projection slender, split distally
and not much differentiated from other 3 processes, a group
of 5-6 strong spines on inner margin (Figure 16). .......
margarsen(p. 25)
This apicodorsal projection broader than any other processes and
with a deep rounded notch apically. ...........200-. 28
Basimere with a long, well developed third subapical projection as
in margursen (Figure 18)... i.) . bos Pees johnsoni (p. 20)
This third subapical projection poorly developed or absent. .....
andamanensis
Basimeres joined subapically, pointed distally and with a small
lobed process on inner margin; distimere expanded at middle
abruptly narrowed and curved distally (Figure 8). .......
fragilis (p. 15)
30
Basimeres separated; distimere varied. .........4.ee68-.
Paraproct bifurcate or divided. 2 6 ec ee Ss ke 31
NOL OS AN0VE. 1925 4 8] Bee 6 i es Oe ge ie ace 33
Paraproct bifurcate distally with 1 prong much shorter than the
other; basimere with a long slender apical projection, a
ventral hairy lobe and 2-pronged dorsal process at base
(Miguve Go 6 a eG a campylostylus(p. 13)
Paraproct divided into 2 large, unequal parts. ..........
Basimere broadly produced distally, terminating into a small, knob-
like process and with a short, pointed subapical inner projec-
MOM LR a ee clavatus
Basimere produced distally into 4 or 5 tooth-like processes, with
many strong bristles and 4 large spines on inner margin. .. .
cautus
Paraproct unusually broad, flattened. ............. . - 04
Paraproct slender,. tapered distally, i... .totoe 35
Distimere with a crescentic process at base and a row of strong
bristles at the middle curved portion; basimere with a group
of strong bristles apically, a 3-pronged process on inner
margin, a long slender process arising from base and a small
basal Beiry lobe. 2 ions Vs ie ele latipennis
Distimere short and markedly swollen distally, with 4 or 5 small
projections and short setae; basimere rather small, simple,
with only a small subapical projection on inner margin. ....
pseudodiurnus
Delfinado: Aedes (Neomacleaya) in Southeast Asia 11
35(33). Basimere with 3 short, differentiated apical and subapical pro-
jections and.a small lobe bearing a group of strong spines;
distimere short and broad, abruptly narrowing to a pointed
tip, with 3-4 long setae at the middle. . .. vallistris
Basimere varied, with 3-4 strong, large, curved spines on inner
Margin... s,s... ee ee ee a Se es 36
36(35). Basimere with forked subapical and apical projections, 3 long
curved spines at base; distimere roughly wedge-shaped in
which one point is narrowly prolonged and bent (Figure 10)...
varietas(p. 40)
Basimere with 2 short projections, one bearing a group of strong
setae, the other toothed, also with a row of 4-6 long, curved
spines on inner margin; distimere curved at middle, narrowed
distally and with scattered long setae. ..... atrius
LARVAE!
Lj Anal gills short, as long as saddle; hairs 9,10,12-P single; 6-I-I
double; 6-III-VI single; 7-I single, branched on other seg-
PCDI 6 iO Se ee as a SS aux
Anal eills longer than saddle. 265 (3.5. io Dea a eS 2
211)... Antennal Deir Poa single see oO ee ed a oe 3
Antennal hair t-A branche@ 2 og 20. ea i I a 5)
3(2). Siphon large and stout, about twice as long as basal width; anal
gills about 2 1/2 times as long as saddle; hair 1-VII with 3-4
short branches; 3-VIII with 5-8 branches (Figure 26); hairs
9,10-P single; hair 1-M branched. .... . Siamensis(p. 37)
Siphon small, slender, about 3 times as long as basal width; anal
gills very long, more than 5 times length of saddle; hair 1-VII
single; 3-VII with at most 4 branches; 9,10-P branched; 1-M
ly arvae of the following unknown: clavatus, cautus, dermajoensis, fragilis,
gibbosus, hispidus, indecorabilis, leicesteri, lugubris, notabilis, neomacro-
dixoa, nubicola, pahangi, panayensis, philippinensis, prioekanensis, pro-
tuberans, pseudodiurnus, varus, singularis, torosus, uncus, vallistris,
varietas and virilis. cyrtolabis and indicus are not included in the key be-
cause there are no specimens available and published descriptions lack de-
finitive characters.
12
4(3).
5(2).
6(5).
15).
8(7).
9(8).
10(9).
11(7).
12(11).
13(12).
14(12).
Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
Thoracic hair 2-M small, single; 9,10,12-P barbed; 3-VIII with
3-4 branches (Figure
PO ee Ss
2-branched (Fieure 90.0 ba ee
. . Yobertsi (p. 36)
Thoracic hair 2-M branched; 9,10, 12-P smooth; 3-VII at most
hamistylus(p. 17)
Abdominal hair 6-II double; 1-P double; 1-A with 4-5 branches;
5,6-C with 2-4 branches. .......... latipennis
Abdominal hair 6-II single; 1-P single; 1-A, 5,6-C at most 2-
branched (Figure d2)icc ork ced k. ishigakiensis (p. 18)
a noreacie lain 6-P doubler) Seer ea 8
Thoracic Dairy @-P siiier ws ee a ewe 11
Thoracic hair 6-T single; 2-P single (Figure 7)
CAG SM, ieee ce Wei Ae Neen DEN eM bk |
campylostylus(p. 14)
Thoracic hair 6-T with 2-4 branches; 2-P branched. ....... 9
Head hair 5, or 6-C with not more than 3 branches, one branch
usually stout and long; abdominal hair 7-I double (Figure 21). .
nigrotarsis (p. 28)
Head hair 5, or 6-C with not less than 6 branches; abdominal hair
PO ea ae Oe aes eee ea. 10
Head hair 8-C double (Figure 18). ......... johnsoni(p. 20)
ats Bair lone and singles. i Oe Oe atrius
Head hair 5-C with not more than 3 branches; 1-P single; 2,3-P
branched (Figure 14). 2S. 257... % - - « macrodixoa (p. 24)
Head hair 5-C with more than 3 branches. ............ 12
Thoracic hairs 9, 12-P Beas SU ee ae ws 13
Thoracic hair 12-P branched; 9-P single. ............ 14
Antennal hair 3-A minute; 6-C with not more than 3 branches; 1-T
Wil 0-6 branches... Bes Bee cretatus
Antennal hair 3-A as long as 6-A; 6-C with more than 3 branches;
1-T with 2-3 weak branches. ...... andamanensis
Head hair 6-C with at most 3 branches; 1-M weak, single. .....
incertus
Head hair 6-C with more than 3 branches; 1-M branched, well
developed (Figure 17)
margarsen (p. 26)
Delfinado: Aedes (Neomacleaya) in Southeast Asia 13
AEDES (NEOMACLEAYA) CAMPYLOSTYLUS LAFFOON
(Figures 6, 7)
Aedes (Aedes) campylostylus Laffoon 1946, J. Wash. Acad. Sci. 36:240
(s*, 9*, L*);1 Knight and Hull 1953, Pacif. Sci. 7:46 (dist. ).
Species with pale lateral patches on the abdominal terga and bare
lower mesepimeron. A. campylostylus can be readily differentiated by the
bifurcate paraproct of the male terminalia and by the characteristic distal
arms of the postatrial plate of the female terminalia. The larva has hairs
1,2-P, 6-T single, 8-P double and abdominal hair 6-I-II double. The pupa
has abdominal hair 2 small but fairly long. It is further characterized by
the following:
FEMALE. Head.Vertex dark brown with light brown narrow scales
around eye margin, patches of pale broad scales laterally and a few up-
right ones mixed with narrow golden scales on occiput; torus with fine hairs
and small flat scales; palpus about 1/8 length of proboscis; proboscis longer
than fore femur by 3 times length of labella, dark brown. Thorax. Scutal
scales dark or reddish brown mixed with golden ones above wing base and
front margin; anterior pronotum with 8-9 bristles and a few narrow scales;
posterior pronotum with 4 bristles, no scales; postspiracular area with 4
bristles, without scales; small patches of pale scales on upper and lower
sternopleuron, 5 bristles behind upper scale patch, without hairs on anterior
portion; upper mesepimeron with a few hairs behind pale scale patch; lower
mesepimeron bare; metameron bare; propleuron with pale scales and 5
bristles. Wing. Alula fringed with brown narrow scales. Legs. Forecoxal
scale patch largely brown with a small pale spot on top; mid and hind coxae
each with pale scales; fore and mid tarsal claws equal, each claw toothed;
hind claws equal, simple. Abdomen. Terga with pale lateral patches;
sterna largely pale scaled, with narrow brown apices. Terminalia.
(Figure 7). Cercus short; postgenital plate broader than long, with a deep
median emargination; postatrial sclerite rounded or nearly so; postatrial
plate with fairly broad, incurved distal arms, hairs confined to median area;
preatrial plate consisting of 2 separated, large, ovate, elongate parts.
Three unequal spermathecae each with a short neck.
MALE. Similar to female in general habitus. Legs. Fore and mid
tarsal claws unequal, only the larger claw toothed; hind claws equal, simple.
Terminalia. (Figure 6). Basimere with a long, slender apical projection,
a ventral hairy lobe and 2-pronged dorsal process at base; distimere curved,
evenly tapered; aedeagus small, rather simple; paraproct expanded at base,
bifurcate distally with 1 prong much shorter than the other.
Lan asterisk (*) following the abbreviations “, ?, P, L indicates that at
least some portion of that sex or stage is figured.
14 Contrib. Amer. Ent. Inst. , vol. 2, no. 4, 1968
LARVA. (Figure 7). Head. Hair 1-C slender; 3-C small, short;
4-C with 4-5 very short branches; 5-C with 6-8 strong branches; 6-C with
6-7 branches; 7-C with 12-16 branches; 8, 10-C with 2-3 branches; 9-C with
4-5 branches; antenna spinose; 1-A inserted at about middle, with 4-5
branches; 2-A longest of the apical hairs; mental plate with 40-42 lateral
teeth. Thorax. Hairs 1,2-P single; 3-P double; 8-P of 2 long branches;
9,10-P single; 12-P double; 1-M 3-branched; 2-M with 2-3 short branches;
3-M single, long; 1-T very small, with 3-4 branches; 2-T with 3-4 fairly
long branches; 3-T small, dendritic tuft; 5-T small, single; 6-T single,
long. Abdomen. Hair 6-I-IL double; 6-DI-VI single; 7-I single, as long as
hair 6, of 4 short branches on II-VI; segment VIII with 12-14 comb scales
arranged in an irregular row, each scale pointed and finely fringed laterally;
hair 1-VIII with 4-5 barbed branches; 2-VIII single or double; 3-VIII with
0-8 barbed branches; 4-VIII double; 5-VIII with 10-12 barbed branches;
Siphon spiculate, with 14-16 long, slender pecten teeth, each tooth with a
lateral denticle except last tooth which is simple; siphonal tuft 1-S 2 or 3-
branched, inserted about 2/3 from base; saddle spiculate; 1-X single; 2-X
with 8-10 branches; 3-X single, very long; anal gills from 2-3 1/2 times
length of saddle, slender and tapered distally; 1-2 precratal tufts.
PUPA. (Figure 6). Cephalothovax. Hairs 1,2,3-C 3-branched;
6-C small, single; 10-C with 6-8 branches; 11-C single, long; 12-C with
3-4 branches. Abdomen. Hair 2-I-VI small, hair-like but a bit longer on
I-II; 3-I double; 3-II-III single; 3-IV with 4-5 branches; 3-V-VII 2-branched;
0-IV-VI of 2 long branches; 5-VII with 2-4 branches; 1-VII of 2 long branches;
6-I-I single, long; 6-III double; 6-IV-VI single, fairly long; 6-VII 2 or 3-
branched; 10-II-IV with 2-3 branches; 10-V-VII single, long; 7-VI-VII
Single; 9-VIM with 2-3 branches, single and poorly developed on other seg-
ments; 4-VII single or double. Paddle. Outer margin finely serrated,
with strong midrib; hair 1-P simple.
TYPE DATA. Holotype male with associated cast skins, from about
4 mi. inland along Balsahan River (near Iwahig), Palawan, PHILIPPINES,
in the U.S. National Museum.
DISTRIBUTION. So far only known from the PHILIPPINES.
Specimens examined: Mindoro; San Jose; Balabac I.; Leyte; Mindanao,
San Ramon, 32 males, 27 females, 12 larvae, 6 pupal skins (USNM, ANS).
TAXONOMIC DISCUSSION. A. campylostylus is strikingly different
in the male terminalia but the female terminalia are in general similar to
those of andamanensis in the development of the postatrial plate; the larva
and pupa are of the same type as andamanensis.
BIOLOGY. Laffoon (1946:242) records that this species prefers
jungle habitats. The larvae have been taken in rock holes, tin cans, gravel-
bottomed residual pools, in a shaded rocky overflow pool and in temporary
road rut pools. Nothing is known of the biting habits.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 15
AEDES (NEOMACLEAYA) FRAGILIS (LEICESTER)
(Figure 8)
Verrallina fragilis Leicester 1908, Cul. Malaya 3:199 (¢’).
Aedes fragilis (Leicester), Edwards 1917, Bull. ent. Res. 7:222 (c'*).
Verrallina malayi Leicester 1908, Cul. Malaya 3:198 (?); Laffoon 1946,
J. Wash. Acad. Sci. 36:238 (taxonomy). NEW SYNONYMY.
Species with pale lateral patches on abdominal terga, toothed hind
tarsal claws, pale scales on the anterior pronotum and bare mesepimeron;
the male and female terminalia are characteristic. The larva and pupa are
not known.
FEMALE. The following description is based on a single female
A. malayi labeled type in the British Museum. Head. Vertex reddish
brown with pale lateral patches, upright scales on occiput not visible; torus
without scales or hairs; palpus about 1/6 length of proboscis; proboscis
about as long as fore femur, brown. Thorax. Scutal scales uniformly
brown; anterior pronotum with some pale broad scales; posterior pronotum
with 4 bristles, without scales; postspiracular area with 4 bristles, without
scales; small patches of pale scales on upper and lower sternopleuron, with-
out hairs on anterior portion; a large pale scale patch on upper mesepimeron,
without hairs behind this scale patch; lower mesepimeron bare; metameron
bare; propleuron with pale scales. Wing. lula fringed with lanceolate
brown scales. Legs. Forecoxal scale patch pale above, brown below; mid
coxa with a small patch of pale scales; a few pale scales on hind coxa; fore
and mid claws equal, each claw toothed; hind claws missing but Leicester
(1908:199) describes "ungues on all the legs equal each with a small tooth".
Abdomen. Terga dark brown with pale lateral patches; sterna with pale
scales. Terminalia. (Figure 8). Cercus long and tapered distally, more
than twice as long as basal width; postgenital plate long and narrow, slightly
emarginate; postatrial sclerite nearly flat at top; postatrial plate fairly
simple, more or less horseshoe-shaped; preatrial plate rather small,
rounded distally. Spermathecae missing.
MALE. Based on the lectotype male A. fragilis in the British
Museum. Externally very similar to the female type specimen labeled
malayi. Legs. Fore legs missing but Leicester (1908:199) describes "fore
and mid ungues unequal, the larger ungues uni-serrate"; mid claws un-
equal, only the larger claw toothed; hind claws equal, each claw toothed.
Terminalia. (Figure 8, after Edwards 1917:222, figure 8i). Basimeres
joined subapically, pointed distally and with a small lobed process on inner
margin; distimere expanded at middle, narrowed and curved distally, with
long hairs; aedeagus appears similar to indecorabilis; paraproct slender,
tapered and curved distally.
: LARVA and PUPA. Unknown.
TYPE DATA. Verrallina fragilis, 1 male and 1 female labeled
cotype in the British Museum. I selected here the male as lectotype with
the following data: "5th mile Gombak Rd, Kuala Lumpur/28/2/04, FED.
16 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
MALAY STATES/G. F. Leicester/1912-350.'' Terminalia and hind claws in
celluloid mount on pin. The female cotype has no abdomen. Vervallina
malayi, a unique female labeled type in the British Museum; terminalia in
celluloid mount on pin; type locality: "Jungle/Gombak Rd 6 mi fr. Kuala
Lumpur, MALAYA!'
DISTRIBUTION: Known only from type locality. Specimens
examined: Kuala Lumpur, 1 male, 2 females (BM).
TAXONOMIC DISCUSSION. Concerning the synonymy of A. malayi
with A. fragilis, Leicester (1908:199, 200) has suggested that malay? is
possibly the female of fragilis or imitator (indecorabilis). A. malayi was
described from a unique female and from Leicester's descriptions and after
examination of the type specimens I have no doubt that the 2 species are con-
specific.
BIOLOGY. Leicester (1908:199, 200) records that the adults have
been caught in the jungle. Larval habitats unknown.
AEDES (NEOMACLEAYA) HAMISTYLUS LAFFOON
(Figures 9, 10)
Aedes (Aedes) hamistylus Laffoon 1946, J. Wash. Acad. Sci. 36:232 (o*, 9*,
L*),
Species with wholly dark abdominal terga and a few pale scales on
anterior pronotum. It is particularly distinctive in having the male fore
tarsal claws, which are toothed, unequal; the mid claws unequal, only the
smaller claw toothed and by the structure of the female postatrial and pre-
atrial plates. The larva can be recognized by the very long anal gills,
single abdominal hair 6-I-VI and single antennal hair 1-A.
FEMALE. Head. Vertex blackish brown with small pale lateral
patches, a few dark upright scales confined to occiput; torus with small, ©
flat dark scales and hairs; palpus about 1/6 length of proboscis; proboscis
about as long as fore femur, dark brown. Thorax. Scutal scales uniformly
dark reddish brown; anterior pronotum with 6 bristles and a few pale scales
which may be rubbed off in some specimens; posterior pronotum with 4-5
bristles, without scales; postspiracular area with 3-4 bristles, without
scales; small loose patches of pale scales on upper and lower sternopleuron,
4 bristles behind upper scale patch, without hairs on anterior portion; upper
mesepimeron with a few hairs behind scale patch; lower mesepimeron bare;
metameron bare; propleuron with pale scales, 2-3 bristles. Wing. Alula
fringed with small flat scales. Legs. Forecoxal scale patch dark brown
with a small pale spot at top; mid and hind coxae each with pale scales; all
tarsal claws equal, each claw toothed. Abdomen. Terga wholly dark brown;
sterna light brown. Terminalia. (Figure 9). Cercus long, tapered distally,
more than twice as long as basal width; postgenital plate rather narrow,
truncate, with a slight median emargination in some specimens; postatrial
sclerite nearly flat at top; postatrial plate quadrate with a small opening, |
without hairs; preatrial plate rounded, small and hairy. Three unequal
spermathecae, without necks.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 17
MALE. Similar to female in general habitus. Legs. Fore tarsal
claws unequal, toothed, the smaller minutely so; mid claws unequal, only the
smaller claw toothed, larger claw simple;hind claws equal, each claw toothed.
Terminalia. (Figure 10). Basimeres joined subapically, roundly produced
distally,bearing a short, blunt process on inner margin; distimere rather
slender, curved distally to a bent, blunt tip; aedeagus long and tapered;
paraproct slender, evenly tapered.
LARVA. (Figure 9). Head. Hair 1-C slender; 3-C very small;
4-C with 3-4 short, weak branches; 5-C usually with 2 strong branches; 6-C
with 2-3 branches; 7-C with 3-4 branches; 8-C with 3-4 fairly long branches;
9-C with 5-6 short branches; 10-C with 2 fairly long branches; antenna spi-
nose; 1-A inserted at about middle of antenna, single; 2-A longest of the api-
cal hairs; mental plate with 34-36 lateral teeth. Thorax. Hairs 1,2-P
single; 3-P with 5-6 short branches; 8-P single; 10-P with 2 long branches;
9,12-P with 4 shorter, smooth branches; 1-M single, long; 2-M with 2-3
short branches; 3-M with 3 long branches; 4-M with 5-6 long branches; 1-T
single, fairly long; 2-T with 3 long branches; 3-T with 6-7 rather long
branches; 4-T with 4 short branches; 5-T small, single; 6-T long, double.
Abdomen. Hair 6-I-VI single; 7-I single, as long as hair 6; 7-II-VI with
3-5 short branches; segment VII with 10-12 comb scales, each scale pointed
and fringed laterally; 1,4-VII single; 2-VIII double, barbed; 3-VIII of 2 long,
barbed branches; 5-VII with 4 barbed branches; siphon small, slender,
about 3 times as long as basal width, spiculate, with 6-7 pecten teeth, each
tooth widely spaced and with 3-5 fine lateral denticles, the last tooth smooth,
inserted from 2/3 to near apex of siphon; siphonal tuft 1-S with 1-3 branches,
subapical; saddle spiculate; 1-X usually single; 2-X with 3-5 branches; 3-X
very long, single; anal gills very long, about 6 times length of saddle; 1-2
precratal tufts.
PUPA. Unknown.
TYPE DATA. Holotype male, Cape Melville, Balabac I.,
PHILIPPINES, in the U. S. National Museum. Paratypes: 41 males, 63
females, 8 larvae, in the U. S. National Museum and British Museum
(Natural History), London.
DISTRIBUTION. Specimens examined: PHILIPPINES: Balabac L.,
Cape Melville, 14 males, 35 females, 4 larvae (USNM); Mindoro, San Jose,
3 males, 2 females (CAS); NORTH BORNEO?: Tawau, 3 males (BM).
TAXONOMIC DISCUSSION. The larva of A. hamistylus is very
similar to robertsi from which it can be differentiated only by minor char-
acters as mentioned in the key. The adults, however, show distinct differ-
ences in the structure of the tarsal claws and terminalia.
BIOLOGY. Laffoon (1946:233) records the larvae were collected in
a gravel-bottomed residual pool in a stream bed; also in a pandanus swamp
by dredging the leaves from the bottom (E; S: Ross).
* New record.
18 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
AEDES (NEOMACLEAYA) ISHIGAKIENSIS BOHART
(Figures 11, 12)
Aedes (Aedes) ishigakiensis Bohart 1956, Bull. Brooklyn ent. Soc. 51:32
ee, eo, Le)
Species with pale lateral patches on abdominal terga, broad pale
scales on anterior pronotum and toothed hind tarsal claws. The male and
female terminalia are particularly distinctive as shown in the figures. The
larva can be distinguished by the single abdominal hair 6-II and thoracic
hair 1-P and branched antennal hair 1-A; the pupa by the long, single ab-
dominal hairs 7,10-VI-VI and branched 6-III-V.
FEMALE. Head. Vertex dark brown with pale lateral patches and
pale scales along eye margin; torus without scales; palpus about 1/6 length
of proboscis; proboscis about as long as fore femur. Thovax. Scutal scales
uniformly brown; anterior pronotum with pale, broad scales; posterior pro-
notum with 3-4 pale scales and 4 bristles; small patches of loose pale scales
on upper and lower sternopleuron, without hairs on anterior portion; post-
spiracular area with bristles; upper mesepimeron with a small patch of pale
scales, reduced or without hairs; lower mesepimeron bare; metameron bare;
propleuron with pale scales. Wing. Alula fringed with brown lanceolate
scales. Legs. Forecoxal scale patch largely brown with pale spot above;
mid and hind coxae with pale scales; tarsal claws of all legs equal, each claw
toothed. Abdomen. Terga with pale lateral patches, that on VII extending
dorsally; sterna with pale basal bands. Terminalia. (Figure 12). Cercus
long, tapered distally, more than twice as long as basal width; postgenital
plate rather narrow, with a shallow median emargination; postatrial sclerite
flat at top; postatrial plate relatively simple, with a pair of sclerotized bars
laterally, large opening and without hairs; preatrial plate markedly large
and expanded laterally, hairy. Three unequal spermathecae without distinct
necks.
MALE.General habitus as in female. Legs. Fore and mid tarsal claws
unequal, both toothed, the smaller minutely so; hind claws equal, each claw
toothed. Terminalia. (Figure 11). Basimeres connected subapically, with
an apical projection bearing a large hooked spine; distimere expanded at
middle, curved distally to a narrow bent tip; aedeagus simple, long and
tapered; paraproct slender, outwardly curved. Tergum IX with a long,
tapered median projection as long as aedeagus.
LARVA. (Figure 12). Based on skins of paratype specimens.
Head. Hair 1-C slender; 3-C very small, single; 4-C with 4-5 short
branches; 5, 6-C of 2 fairly strong branches; 7-C with 5-7 branches; 8,10-C
with 3-4 branches; 9-C with 4-6 branches; antenna spinose; 1-A inserted at
about middle of antenna, 2-branched; 2-A longest of the apical hairs; mental
plate with 28-32 lateral teeth. Thorax. 1-P single; 2,3-P with 2-3 branches;
8,10,12-P single; 9-P double, shorter than 10,12-P; 1-M with 3 branches; :
2-M very small, branched; 3-M with 2 weak branches; 4-M with at most 6
branches; 1-T with 4 short branches; 3-T with 8-10 branches; 4-T with 3-5
Delfinado: Aedes (Neomacleaya) in Southeast Asia 19
short branches; 5-T small, single; 6-T double, fairly long. Abdomen.
Hair 6-I 2-3 branched; 6-II-VI single; 7-I single; 7-II-VI with 3-4 short
branches; segment VIII with 10-12 comb scales, each scale pointed and with
lateral fringe; hair 1-VIII with 4-5 barbed branches; 2-VII with 2-3 weak
branches; 3-VIII with 5-7 long barbed branches; 4-VIII single; 5-VIII with
6-8 barbed branches; siphon rugose, about 2 1/2 times as long as basal
width, with 11-13 pecten teeth, each tooth with a lateral denticle, some-
times last distal tooth simple and inserted at about middle of siphon; si-
phonal tuft 1-S with 3-4 branches, inserted beyond middle of siphon; saddle
rugose; 1-X double; 2-X with 3-4 branches; 3-X long, single; anal gills
about twice as long as saddle; 2 precratal tufts.
PUPA. (Figure 11). Description based on paratype specimens.
Cephalothorax. Hairs 1,2-C bifid; 3-C single; 6-C small, double; 10-C
with 5 branches; 11-C single; 12-C with 4 branches. Abdomen. Hair
2-II-VI strong, almost spine-like; 3-I with 3 branches; 3-II-III long, Single;
3-IV-VI with 3-5 short branches; 5-IV short, double; 5-V-VI very long,
single; 4,5-VII with 3-4 branches, subequal in length; 6-I bifid or trifid;
6-II very long, single; 6-III-VII with 2-4 branches; 7,10-VI-VI single, long;
10-II-IV with 3 short branches; 9-VII with 2 short branches, very short
and single on other segments; 4-VIII with 2-3 weak branches. Paddle.
Outer margin serrated; hair 1-P simple.
TYPE DATA. Holotype male with associated cast skins, Mt.
Banna, Ishigaki, RYUKYUI., in the U. S. National Museum.
DISTRIBUTION. Known only from type locality. Specimens exam-
ined: RYUKYU I.: Mt. Banna, Ishigaki, 1 male, 14 females (USNM).
TAXONOMIC DISCUSSION. A. ishigakiensis shows close relation-
ship with fragilis, hamistylus, indecorabilis and siamensis on the basis of
male and female terminalia. The presence of pale lateral markings on ab-
dominal terga differentiates it from the above mentioned species, and from
all other species by the presence of a long, tapered median projection of
male tergum IX.
BIOLOGY. | Bohart (1956:33) records the larvae have been taken
from a foxhole in pine woods. Nothing is known of the biting habits.
AEDES (NEOMACLEAYA) JOHNSONI LAFFOON
(Figures 18, 19)
Aedes (Aedes) johnsoni Laffoon 1946, J. Wash. Acad. Sci. 36:235 (o*, 9*, L).
The adults of this species are very similar to A. margarsen and A.
andamanensis by having fine hairs on metameron, forked apical projections
on male basimere and by the characteristic distal arms of the female post-
atrial plate. The male, however, can be recognized by the much broader
apicodorsal projection of the basimere, also it has a long, well developed
third subapical projection; the female by the widely open distal arms of the
20 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
%
postatrial plate. The larva has thoracic hair 8-P double, and hair 6-I-V
long and double in the pupa.
FEMALE. Head. Vertex dark brown with pale lateral patches,
some narrow golden scales around eye margin and upright forked scales on
occiput; torus with small narrow dark scales. Thorax. Scutal scales golden
brown, paler on anterior margin and above wing base; anterior pronotum
with about 12 bristles, without scales; posterior pronotum with some pale
brown narrow scales and bristles; postspiracular area with bristles and
scales; a large patch of pale scales on upper sternopleuron, without hairs
on anterior portion; upper mesepimeron with hairs behind pale scale patch
extending downwards to middle of sclerite; lower mesepimeron bare; meta-
meron with fine hairs; propleuron with pale scales. Wing. Alula fringed
with narrow brown scales. Legs. Fore and mid tarsal claws equal, toothed;
hind claws equal, simple. Abdomen. Terga with pale lateral patches pro-
duced on to dorsum; sterna pale basally, brown apically except VII which
appears mostly brown. Terminalia. (Figure 19). Very similar to
margarsen. Distal arms of postatrial plate widely open and gently curved
while in margarsen they are straight and directed inwards. One large and
2 small, subequal spermathecae each with a short neck.
MALE. General habitus as in female. Legs. Fore and mid tarsal
claws unequal, only the larger claw toothed; hind claws equal, simple.
Terminalia. (Figure 18). Very similar to margarsen or andamanensis ex-
cept as noted: the apicodorsal projection of the basimere is much broader
than any other processes and it has a deep notch apically; this projection
narrower and split distally in margarsen; also it has a long, well developed
third subapical projection as in margarsen but absent or poorly developed in
andamanensis . |
LARVA. (Figure 18). Head, 1-C slender; 3-C very small, single;
4-C with 8 very short branches; 5, 6-C with 6-8 barbed branches; 7-C with
12-14 barbed branches; 8-C usually double; 9,10-C with 2-5 shorter branches;
antenna spinose; 1-A inserted a little below middle, with 4-5 branches; 2-A
longest of the apical hairs; mental plate with 28-32 lateral teeth. Thorax.
1-P single; 2,3-P with 2-3 branches; 8-P 2-branched, long; 9, 10-P single;
12-P double; 1-M with 3-4 short branches; 2-M very small, with 5 branches;
3-M single, very long; 1-T with 4-5 short branches; 2-T with 4-5 long
branches; 3-T with many short branches; 5-T small, double; 6-T double,
long. Abdomen. Hair 6-I-II double; 6-III-VI single; 7-I single, rarely
double, branched on other segments; segment VIII with 10-14 comb scales,
each scale pointed and with fine lateral fringe; hair 1-VIII with 4-7 barbed
branches; 2, 4-VIII double; 3-VIII with 3-6 barbed branches; 5-VIII with 6-9
barbed branches; siphon with 14-20 pecten teeth, last tooth inserted beyond
middle, basal teeth with small lateral denticles, distal 2-5 teeth simple or
with a small denticle, more widely spaced than those nearer the base; si-
phonal tuft 1-S inserted about 2/3 from base, with 2-5 branches; saddle spi-
culate; 1-X single; 2-X with 5-7 branches; 3-X single; anal gills 2-3 times
length of saddle; 2 precratal tufts.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 21
PUPA. Chaetotaxy as in figure 19. Cephalothorax. Hairs
1,2,3-C 2-3 branched; 6-C single; 10-C with 6-8 branches; 11-C single or
bifid; 12-C with 3-5 branches. Abdomen. Hair 2-I-VII small, hair-like but
slightly longer on I-II; 3-I with 3-4 branches; 3-II-III single, long; 5-IV-VI
double; 5-VII usually single; 6-I-V double, long; 6-VI single; 6-VII with 4-6
branches; 7,10-VI-VII single; 9-VIII with 2-3 branches; 9-VII single, long
but poorly developed on other segments. Paddle. Outer margin finely ser-
rated, with fairly strong midrib; hair 1-P simple.
TYPE DATA. Holotype male with associated cast skins, San Ramon,
Pasonanca, Zamboanga, Mindanao, PHILIPPINES, in the U. S. National
Museum.
DISTRIBUTION. Apparently restricted to Mindanao, PHILIPPINES.
Specimens examined: Kabakan, 35 females, 4 males; Zamboanga, Pasonanca
and San Ramon, 31 females, 24 males, 30 pupal and larval cast skins;
Basilan I., Isabela, 4 males, 5 females (USNM).
BIOLOGY. Laffoon (1946:236) records that the larvae have been
collected in muddy pools and road ruts. Adults have been taken in the jungle
during the daytime.
AEDES (NEOMACLEAYA) LEICESTERI EDWARDS
(Figure 8)
Aedes leicesteri Edwards 1917, Bull. ent. Res. 7:222 (o*); Barraud 1934,
Fauna Brit. India, Dipt. 5:291.
Species with fine hairs on the lower mesepimeron and pale lateral
patches on the abdominal terga. The male aedeagus is very similar to those
of uncus, protuberans or torosus in being very large but differs by having a
much enlarged distimere; also the basimere has a large, curved spine on
the inner margin and a short subapical projection armed with 2 large and 3
small spines. The female, larva and pupa are not known.
MALE. Based on a single poor specimen from North Borneo and a
unique type in the British Museum. Head. Vertex mostly brown with some
pale scales laterally, a few upright scales on occiput; palpus about 1/8
length of proboscis; proboscis longer than fore femur. Thovax. Scutal
scales brown; most of the pleural scales rubbed off but there is a patch of
pale scales on upper mesepimeron, and fine hairs behind these scales ex-
tend downwards to lower portion; metameron bare. Wing. Alula fringed
with narrow scales. Legs. Fore and mid tarsal claws unequal, only the
larger claw toothed; hind claws gone in both specimens. Abdomen. Terga
with pale lateral patches; sterna brown with scattered pale scales. Termi-
nalia. (Figure 8). Basimere slightly projected apically with a large curved
spine on inner margin and a short subapical projection armed with 2 large
and 3 small spines; distimere much enlarged and broadened distally, witha
few long setae; aedeagus very large, of uncus type; paraproct very short,
pointed distally.
22 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
FEMALE, LARVA and PUPA. Unknown.
TYPE DATA. A unique male labeled type in the British Museum
(Natural History), terminalia in celluloid mount on pin, Ampang Jungle,
Kuala Lumpur, MALAYA. :
DISTRIBUTION... Specimens examined: MALAYA: Kuala Lumpur,
1 male (BM); NORTH BORNEO: N. of Kalabakan, 1 male (Bishop).
BIOLOGY. Unknown.
AEDES (NEOMACLEAYA) LUGUBRIS BARRAUD
(Figure 13)
Aedes (Aedes) lugubris Barraud 1928, Indian J. med. Res. 16:3'72 es
Barraud 1934, Fauna Brit. India, Dipt. 5:294 (¢*).
A. lugubris is strikingly different from other species by having only
one complete pale band on abdominal terga, by the fairly simple female ter-
minalia and by the horn-like apical projection of the male basimere as shown
in the figure. The larva and pupa are not known.
FEMALE. Head. Vertex blackish brown with a line of pale scales
at middle, patches of pale scales laterally and a few upright ones confined
to occiput which are rather difficult to see; torus with a few fine hairs; pal-
pus about 1/6 length of proboscis; proboscis longer than fore femur by length
of labella, dark brown. Thorax. Scutal scales uniformly brown to reddish
brown; anterior pronotum with 8 bristles, a few flat scales and fine hairs;
posterior pronotum with some narrow brown scales and 5 bristles; post-
spiracular area with 5 bristles and a few scales; upper sternopleuron with a
large patch of pale scales, a smaller patch below, without hairs on anterior
portion; upper mesepimeron with a patch of pale scales, hairs confined to
area behind this patch; lower mesepimeron bare; metameron bare; pro-
pleuron with a fairly large patch of pale scales. Wing. Alula fringed with
lanceolate scales, a few appearing broad. Legs. Forecoxal scale patch
largely brown with a small pale spot above; mid and hind coxae each with a
row of pale scales; fore and mid claws unequal, each claw toothed; hind claws
equal, simple. Abdomen. Terga with a complete pale band only on tergum
II, pale lateral patches on other segments; sterna largely pale with brown
apices. Terminalia. (Figure 13). Cercus rather short, about twice as long
as basal width; postgenital plate large, with a shallow median emargination;
postatrial sclerite convex; postatrial plate fairly simple, hairy below the
opening; preatrial plate poorly sclerotized, joined at one end, hairy. Three
unequal spermathecae each with a neck.
MALE. General habitus as in female. Legs. Fore and mid tarsal
claws unequal, only the larger claw toothed; hind claws unequal, simple.
Terminalia. (Figure 13). Basimere with a horn-like apical projection, 2
shorter, simple subapical processes and a bifurcate median process; disti-
mere curved and pointed distally, with a small subapical process; aedeagus
small, very simple; paraproct poorly developed.
|
Delfinado: Aedes (Neomacleaya) in Southeast Asia 23
LARVA and PUPA. Unknown.
TYPE DATA. I have seen the type female in the British Museum
labeled: "type/Rangoon, BURMA/20/v/26, Capt. R. W. Hamilton-Miller. "
Also another female on a slide labeled cotype with same locality and date as
type. |
DISTRIBUTION. Specimens examined: MALAYA: Selangor, 8
females, 2 males (USNM), 3 females (Bishop); BURMA: Rangoon, 2 females
(BM). This species has also been reported from the ANDAMAN I., but I
have not seen specimens from this locality.
BIOLOGY. Unknown.
AEDES (NEOMACLEAYA) MACRODIXQA DYAR AND SHANNON
(Figures 14, 15)
Aedes (Aedes) macrodixoa Dyar and Shannon 1925, Insec. Inscit. menst.
13:79 (o); Laffoon 1946, J. Wash. Acad. Sci. 36:233 (“*, 9*).
Species with fine hairs on the lower mesepimeron and pale lateral
patches on the abdomen which are slightly produced on to dorsum. The male
is particularly distinctive by the markedly developed Y-shaped structure on
tergum IX, and the female by the shape of the spermathecae. The larva has
head hairs 5, 6-C with not more than 3 branches; thoracic hairs 8-P, 3-M
single and abdominal hair 6-I-IJ double. The pupa is not known.
FEMALE. Head. Vertex dark brown with a line of narrow golden
scales at middle, pale patches of broad scales laterally, a few upright scales
confined to occiput; torus with a few hairs and flat scales; palpus about 1/8
length of proboscis; proboscis longer than fore femur by twice length of
labella, dark brown. Thovax. Scutal scales dark brown with some golden
ones on front margin, prescutellar area, scutellum and above wing base;
anterior pronotum with about 10 bristles, without scales; posterior pronotum
with 6 bristles and a few narrow scales; postspiracular area with 5-6
bristles, without scales; sternopleuron with a large pale scale patch on upper
portion and 7 bristles behind this patch, a smaller patch of pale scales mixed
with fine hairs below, anterior portion with many fine hairs; upper mesepi-
meron with a patch of pale scales, hairs behind this patch extend downwards
to lower portion of sclerite; metameron bare; propleuron with a small patch
of pale scales. Wing. Alula fringed with broad scales. Legs. Forecoxal
scale patch dark brown with a pale spot above; mid and hind coxae each with
patches of pale scales; fore and mid claws equal, each claw toothed; hind
claws equal, simple. Abdomen. Terga with pale lateral patches slightly
produced on to dorsum; sterna pale-scaled at basal 2/3, brown apically.
Terminalia. (Figure 14). Cercus rather short, about twice as long as basal
width; postgenital plate broader than long, with a shallow median emargina-
tion; postatrial sclerite rounded or nearly so; postatrial plate with lateral
expansions, a few short hairs around opening; preatrial plate large, faintly
joined at both ends, with wrinkled margin. Three unequal spermathecae
each with a swollen neck.
24 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
MALE, Similar to female in general habitus. Legs. Fore and
mid tarsal claws unequal, only the larger claw toothed; hind claws equal,
simple. Tervminalia. (Figure 15). Basimere with a long, slender projec-
tion between 2 short ones and a large, pointed median projection on inner
margin; distimere fairly slender for most of its length, then curved distally;
aedeagus large, simple, of the wncus type; paraproct very small, poorly
developed. Tergum IX with a markedly large median Y-shaped structure
similar to neomacrodixoa but the branches of the Y-shaped structure are
wider and more abruptly tapered and without lateral lobe.
LARVA. (Figure 14). Based on cast skins. Head. 1-C slender;
3-C short; 4-C with 4-5 short branches; 5-C with 2-3 strong branches; 6-C
with 1-3 strong branches, usually single in most specimens; 7-C with 6-8
strong branches; 8, 9,10-C with 3-4 short branches; antenna spinose; 1-A
inserted near middle, with 4-6 branches; 2-A longest of the apical hairs;
mental plate with about 34 lateral teeth. Thovax. 1-P single; 2,3-P either
2 or 3-branched; 8-P single; 9,10-P single; 12-P double; 1-M with 3-4 short
branches; 2-M very small, branched; 3-M single, fairly long; 1-T with 4
short branches; 2-T with 5-6 long branches; 3,4-T very small, branched;
5-T small, single; 6-T 2 or 3-branched. Abdomen. 6-I-II double; 6-III-VI
single; 7-I single; 7-II-VI with 3-5 short branches; segment VIII with 12
pointed comb scales, each scale with fine lateral fringe; 1-VIII with 4-5
barbed branches; 2-VIII with 2-3 weak branches; 3, 5-VIII with 6-10 strong,
barbed branches; 4-VIII usually 2-branched; siphon rugose, with 16-18
pecten teeth, each with 1-4 denticles, distal 3-4 teeth more widely spaced
than those nearer the base; siphonal tuft 1-S inserted 2/3 from base, with
2-4 branches; saddle rugose; 1-X single; 2-X with 7-9 branches; 3-X single,
long; anal gills about 3 times length of saddle; 2-3 precratal tufts.
PUPA. Unknown.
TYPE DATA. Holotype male, Infanta, (Tayabas), Quezon, Luzon,
PHILIPPINES, in the U. S. National Museum.
DISTRIBUTION. Specimens examined: PHILIPPINES: Mindoro,
San Jose, 26 males, 17 females; Leyte, Tacloban, 2 males, 1 female;
Mindanao, Kabakan, 8 males, 16 females (USNM).
TAXONOMIC DISCUSSION. This species is indistinguishable ex-
ternally from uncus, nubicola or neomacrodixoa. The male terminalia
closely resemble those of neomacrodixoa but the hairy branch of the Y-shaped
structure of tergum IX of neomacrodixoa is expanded and with a lateral lobe,
whereas it is tapered and without such a lobe in macrodixoa.
BIOLOGY. Laffoon (1946:239) records that the adults were bred
from larvae taken in temporary leaf-filled pools; others were captured in
dense jungle during the daytime. Larvae have also been collected in shaded
temporary puddles and hoof prints (E: S. Ross). Nothing is known of the
biting habits.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 25
AEDES (NEOMACLEAYA) MARGARSEN DYAR AND SHANNON
(Figures 16, 17)
Aedes (Aedes) margarsen Dyar and Shannon 1925, Insec. Inscit. menst.
13:80 (c, 2); Bohart 1945, U. S. Navmed 580, figs. 51, 57 (a, gf);
Laffoon 1946, J. Wash. Acad. Sci. 36:237 (c*, L).
Species with fine hairs on metameron, pale lateral patches on ab-
dominal terga, forked apical projections on male basimere and characteris-
tic distal arms of female postatrial plate. The male is recognized by the
slender apicodorsal projection split distally and not much differentiated from
other processes, also by the presence of a long, well developed third sub-
apical projection on the basimere. The larva has hairs 1,8-P single, 3-M
long and single and 1-M branched and well developed. The pupa is most
difficult to separate from campylostylus; it has hair 6-I-II single; 6-I-V
with 2-3 branches and 7,10-VI-VII single.
FEMALE. Vertex dark brown with narrow pale scales around eye
margin, middle and occiput, patches of pale scales laterally, a few upright
ones confined to occiput; torus with fine hairs and small flat scales; palpus
about 1/6 length of proboscis; proboscis about as long as fore femur, brown.
Thorax. Scutal scales brown with golden ones around front margin and pre-
scutellar area; anterior pronotum with about 10 bristles, a few pale scales;
posterior pronotum with narrow golden scales, 6 bristles; postspiracular
area with 3-4 bristles, a few scales; upper sternopleuron with a large patch
of pale scales, a row of 7 bristles behind this patch, without hairs on ante-
rior portion; upper mesepimeron with hairs behind pale scale patch extending
downwards to middle of sclerite; lower mesepimeron bare; metameron with
many fine hairs; propleuron with a large patch of pale scales. Wing. Alula
fringed with narrow brown scales. Legs. Forecoxal scale patch largely
brown; mid and hind coxae each with patches of pale scales; fore and mid
claws unequal, each claw toothed; hind claws equal, simple. Abdomen.
Terga with pale lateral patches produced on to dorsum, occasionally form-
ing complete bands in some specimens; sterna pale-scaled with brown apices.
Terminalia. (Figure 17). Very similar to johnsoni except that the distal
arms of postatrial plate are straight and directed inwards, the postgenital
plate has narrower emargination and the preatrial plate consists of 2 rectan-
gular parts slightly joined at middle. One large and 2 small subequal sper-
mathecae each with a short neck.
MALE. General habitus as in female. Legs. Fore and mid claws
unequal, only the larger claw toothed; hind claws equal, simple. Terminalia.
(Figure 16). Similar to those of johnsoni or andamanensis but basimere
with slender apicodorsal projection, split distally and not much differentiated
from other 3 processes, a third long subapical projection as in johnsoni,
and a group of 5-6 stronger spines on inner margin; distimere slender for
most of its length and curved beyond middle; paraproct long, swollen at
base, evenly tapered as in the above mentioned species.
26 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
LARVA. (Figure 17). Head. 1-C slender; 3-C short; 4-C with
6-8 very short branches; 5, 6-C with 6-9 branches; 7-C with at most 14
branches; 8-C usually with 2 short branches; 9,10-C with 3 branches; an-
tenna spiculate; 1-A inserted before middle, with 5 branches; 2-A longest
of the apical hairs; mental plate with 28-34 lateral teeth. Thorax. 1,8-P
single, long; 2-P normally 2-branched; 3-P with 3-4 branches; 9, 10-P
single; 12-P 2 or 3-branched; 1-M with 4-5 long branches; 2-M with 3 short
weak branches; 3-M long, single; 4~M with 4 long branches; 1-T small, with
9 Short weak branches; 2-T with 4-5 long branches; 3-T small tuft with many
branches; 4-T very small branched tuft; 5-T small, single; 6-T with 3 long
branches. Abdomen. 6-I-II double; 6-III-VI single; 7-I single; 7-II-VI
branched, decreasing in length posteriorly; segment VIII with 9-11 comb
scales, each scale pointed and with fine lateral fringe; 1-VIII with 4-5
branches; 2, 4-VII 2-branched; 3-VIII with 5-6 longer branches; 5-VII with
7-9 branches; siphon rather smooth, with 14-16 pecten teeth, distal 3-4
teeth widely spaced and simple, basal teeth each with a lateral denticle;
siphonal tuft 1-S inserted about 2/3 from base, with 2-3 branches; saddle
finely rugose; 1-X single; 2-X with 5-7 branches; 3-X long, single; anal
gills about twice length of saddle; 1-2 precratal tufts.
PUPA. (Figure 16). Cephalothorax. Hairs 1,2,3-C with 2-3
branches; 6-C 2-branched; 10-C with 9-12 branches; 11-C single; 12-C with
3-5 branches. Abdomen. 2-I-VII small, hair-like but slightly longer on
I-IT; 3-I, IV-VII with 3-5 branches; 3-II-III single, long; 5-II-IV with 3-5
branches; 5-V-VI with 2 long branches; 6-I-II, VI single, long; 6-III-V, VI
with 2-4 branches; 7,10-VI-VII single; 9-VIII with 2-3 branches, occasionally
4-branched in some specimens; 9-VII with 1-2 branches; single and poorly
developed on other segments. Paddle. With finely serrated margin, strong
midrib; 1-P simple.
TYPE DATA. Lectotype male (selected by Laffoon 1946:237),
Camp Eldridge, Laguna, Luzon, PHILIPPINES, in the U. S. National
Museum, terminalia on slide.
DISTRIBUTION. Specimens examined; PHILIPPINES: Mindoro,
San Jose, 41 males, 45 females, 10 larvae, 2 pupal skins (CAS); Luzon, La
Union, Anastacio, 10 males; Laguna, Camp Eldridge, 3 males, 4 females;
Cagayan, 1 female; Zambales, Subic Bay, 1 male, 3 females (USNM, JHS).
TAXONOMIC DISCUSSION. A. margarsen is apparently closely
related to johnsoni and andamanensis on the basis of external characters
and terminalia as well as on certain chaetotaxy of the immature stages.
However, differences in the terminalia of the 3 species are quite distinctive
as discussed elsewhere.
BIOLOGY. Laffoon (1946:237) records the larvae have been taken
in road rut pools. Nothing is known of the biting habits.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 27
_ AEDES (NEOMACLEAYA) NEOMACRODIXOA KING AND HOOGSTRAAL
(Figure 15)
Aedes (Aedes) neomacrodixoa King and Hoogstraal 1947, J. Wash. Acad.
Sci. 37:124 (o*, ¢). .
The adults of this species are indistinguishable from those of uncus,
nubicola or macrodixoa. The line of pale scales on the occiput which King
and Hoogstraal regarded as being diagnostic for this species also occurs in
macrodixoa. The terminalia are very similar to those of macrodixoa but
can be recognized by the Y-shaped structure of tergum IX and by the pres-
ence of 1-2 rod-like setae on outer apical projection of basimere. The larva
and pupa are not known.
FEMALE. Head. Vertex dark with pale lateral patches, narrow
golden ones mixed with upright scales on occiput extending forward to be-
tween the eyes. Thovax. Scutal scales largely brown, golden anteriorly,
over wing base and prescutellar area; anterior pronotum without scales;
posterior pronotum with scattered narrow brown scales and about 6 bristles;
upper sternopleuron with a large patch of pale scales and fine hairs on ante-
rior portion; mesepimeron with numerous fine hairs behind pale scale patch
and lower part of sclerite; metameron bare. Legs. Fore and mid tarsal
claws equal, each claw toothed; hind claws equal, simple. Abdomen. Terga
with pale lateral patches well produced on to dorsum; sterna largely pale
basally. Terminalia. (Figure 15). Essentially similar to those of
macvodixoa. I find no characters for separating the females of these except
that in neomacrodixoa the postgenital plate is deeply emarginate and usually
only the largest spermatheca has a large, swollen neck.
MALE. General habitus as in female. Legs. Fore and mid tarsal
claws unequal, only the larger claw toothed; hind claws equal, simple.
Terminalia. (Figure 15). Very similar to those of macrodixoa except as
follows: basimere with, 1-2 rod-like setae on outer apical projection, (these
are weak in macrodixoa,)numerous strong setae dorsally and a more pro-
nounced inner apical projection; the Y-shaped structure of tergum IX has a
longer stem, more hairy apical branches, each branch expanded and with a
lateral lobe.
LARVA and PUPA. Unknown.
TYPE DATA. Holotype male, Hollandia, NETHERLANDS NEW
GUINEA, in the U. S. National Museum.
DISTRIBUTION. Specimens examined: NEW GUINEA: 19 males,
17 females (paratype series, USNM); CELEBES: Kuluwara, 1 male
(Bishop).
BIOLOGY. Adults have been collected at light and Malaise traps.
Nothing is known of the breeding habitats.
28 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
AEDES (NEOMACLEAYA) NIGROTARSIS (LUDLOW)
(Figures 20, 21)
Pseudoskusea nigrotarsis Ludlow 1908, Canad. Ent. 40:52 (9).
Aedes (Aedes) nigrotarsis (Ludlow), Laffoon 1946, J. Wash. Acad. Sci.
36:239 (o*, 9*, L).
Species with complete pale bands on abdominal terga. The male
terminalia are particularly distinctive in having a large rectangular apical
projection and a crescentic process near the base of basimere; the female |
by the darkly pigmented preatrial plate and by the spermathecae each with
a large bulbous neck. The larva has 8-P double; 1-P, 3-M single and ab-
dominal hair 7-I double; the pupa has hairs 1,3,5-IV-VII with 3 or more
branches and 6-IJ-VI with 3-4 branches.
FEMALE. Head. Vertex blackish brown with some narrow golden
scales on occiput, broad pale scales laterally and upright ones confined to
occiput sometimes difficult to see; torus with small flat scales and a few
hairs; palpus about 1/6 length of proboscis; proboscis longer than fore femur
by length of labella. Thorax. Scutal scales deep brown with some golden
ones on front margin, above wing base, prescutellar area and scutellum;
anterior pronotum with about 9 bristles and short setae, a few scales pres-
ent on some specimens; posterior pronotum with narrow brown scales and 4
bristles; postspiracular with 4 bristles and narrow curved golden scales;
patches of loose scales on sternopleuron and upper mesepimeron; 5-6 bris-
tles behind pale scales on upper sternopleuron, without hairs on anterior
portion; a few hairs behind pale scales on upper mesepimeron; lower mes-
epimeron bare; metameron bare; a few loose scales on propleuron. Wing.
Alula fringed with dark brown lanceolate scales. Legs. Forecoxal scale
patch largely brown; mid and hind coxae each with pale scales; fore and mid
tarsal claws equal, each claw toothed; hind claws equal, simple. Abdomen.
Terga with complete pale subbasal bands which may be broken dorsally in
some specimens; sterna largely dark with some scattered pale scales.
Terminalia. (Figure 21). Cercus tapered distally, about twice as long as
basal width; postgenital plate slightly longer than wide, with a fairly deep
median emargination; postatrial sclerite rounded or nearly so; postatrial
plate with a fairly large cordate opening and thick long hairs; preatrial plate
entire, darkly pigmented. Three unequal spermathecae each with a large
bulbous neck and punctate in this region.
MALE. Similar to female in general habitus. Legs. Fore and mid
tarsal claws unequal, only the larger claw toothed; hind claws equal, simple.
Terminalia. (Figure 20). Basimere with a large rectangular apical projec-
tion and a crescentic process near base; distimere slender, with a small
thumb-like process distally; aedeagus rather small, of the andamanensis
type; paraproct expanded at base, tapered distally and curved inwards.
LARVA. (Figure 21). Based on cast skins. Head. 1-C fairly
slender; 3-C rather long; 4-C with 5-6 very short branches; 5, 6-C with not
more than 3 branches, one branch usually stout and long if 3-branched; 7-C
Delfinado: Aedes (Neomacleaya) in Southeast Asia 29
with 8-10 branches; 8-C with 3 long branches; 9,10-C with 3-4 short
branches; antenna spinose; 1-A inserted before middle, with 4-5 branches;
2-A longest of the apical hairs; mental plate with about 40 lateral teeth.
Thorax. 1-P usually single; 2-P branched apically; 3-P with 3-4 branches;
8-P double; 10, 12-P single; 9-P with 3 short branches; 1-M with 4-5 fairly
long branches; 2-M very small, branched; 3-M single, long; 1-T very small,
branched; 2-T with 5-7 branches; 3-T many-branched short tuft; 4-T very
small branched tuft; 6-T with 4-5 long branches. Abdomen. 6-I-II with 2-3
branches; 6-III-VI single; 7-I double; 7-I with 5 strong barbed branches;
7-III- VI with 6-8 weak short branches; segment VIII with 10 pointed comb
scales, each scale with fine lateral fringe; 1-VIII with 3-5 barbed branches;
2-VIlI with 3-4 branches; 3-VIII with 8-12 barbed branches; 4-VIII single;
o-VIII with 6-8 barbed branches; siphon rugose, rather stout, about twice as
long as basal width; with 10-11 pecten teeth, each tooth with 1-2 lateral den-
ticles, distal 2 teeth widely spaced, last tooth simple and inserted a little
beyond middle; siphonal tuft 1-S with 3-5 weak branches, inserted 2/3 from
base; saddle rugose, broader than long; 1-X single; 2-X with 10-12 branches;
3-X single; anal gills about 2 1/2 times length of saddle; 2 precratal tufts.
PUPA. (Figure 20). Cephalothorax. 1,2,3-C with 2-3 branches;
6-C short, single; 10-C with 5-7 branches; 11-C single; 12-C with 3-4
branches. Abdomen. Setae mostly short, branched. 2-I-VII small, hair-
like except on I, II where it is stronger; 3-III long, single; 3-I-II, IV-VII
with 2-6 branches, comparatively longer on V-VI; 5-IV-VII with 3-5 long
branches; 6-I-II single, long; 6-III-VI with 3-4 long branches; 6-VII short,
branched; 7-VI, 10-VI-VII double; 7-VII single; 9-VIII 2-branched, occa-
sionally triple; 9- VII with 1-3 much shorter branches, single and inconspic-
uous on other segments. Paddle. With finely serrated outer margin; 1-P
simple.
TYPE DATA. Holotype female, Infanta, (Tayabas) Quezon, Luzon,
PHILIPPINES, in the U. S. National Museum.
DISTRIBUTION. Specimens examined: PHILIPPINES: Luzon, La
Union, 7 females; Rizal, 16 females, 9 males (USNM); Cagayan, 11 females,
2 males (JHS); Leyte, 27 females, 5 males (USNM, Bishop); Calicoan I. ,
1 male; Samar, 5 males, 2 females with cast skins (ANS); Mindanao, 30 fe-
males, 2 males, 15 pupal and larval skins (JHS), 2 males (CNHM).
TAXONOMIC DISCUSSION. A. nigrotarsis is apparently closely
related to indicus. The male terminalia, however, differ from indicus by
the process near the base of the basimere which is crescentic, not long and
slender as figured for indicus by Barraud (1934:284). Also, I have examined
the female type of indicus at the British Museum and I found significant dif-
ferences in the details of the terminalia and shape of spermathecae. The
preatrial plate consists of 2 darkly pigmented separate parts in indicus,
whereas it is entire in nigrotarsis. 3
BIOLOGY. Laffoon (1946:240) records that the larvae have been
collected several times in temporary grassy pools, and the adults at light
traps. Nothing is known of the biting habits.
30 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
AEDES (NEOMACLEAYA) NUBICOLA LAFFOON
(Figure 22)
Aedes (Aedes) nubicolus Laffoon 1946, J. Wash. Acad. Sci. 36:237 (o*);
Bohart 1945, U. S. Navmed 580, fig. 50 (as wncus).
Species with fine hairs on lower mesepimeron and pale lateral
patches on abdominal terga. The adult male can be recognized by the
characters of the terminalia, namely, the presence of 5 large distal spines
on apical projection of basimere, the short and broad distimere and by the
very long slender paraprocts which have a pair of elongate structures a-
rising between them. The female, larva and pupa are not known. The fol-
lowing description is based on a single male paratype.
MALE. Head. Vertex dark brown with scattered pale scales later-
ally and around eye margin, many upright ones on occiput; torus bare; palpus
about 1/8 length of proboscis; proboscis longer than fore femur by twice
length of labella, dark brown. Thorax. Scutal scales dark brown, paler at
front margin; anterior pronotum with 7 bristles, without scales; posterior
pronotum with a few dark narrow scales, 3 bristles; spiracular area with
6 bristles, without scales; sternopleuron and upper mesepimeron with few
pale scales; anterior portion of sternopleuron with fine hairs; many fine
hairs behind pale scales on upper mesepimeron which extend downwards to
lower portion of sclerite; metameron bare; propleuron with a few pale scales.
Wing. Alula fringed with flat scales. Legs. Forecoxal scale patch dark
brown with pale scales on top and bottom; mid coxa with a small patch of
pale scales; none on hind coxa; fore and mid tarsal claws unequal, only the
larger claw toothed; hind claws equal, simple. Abdomen. Terga with pale
lateral patches; sterna mostly with scattered pale scales. Terminalia.
(Figure 22), Basimere with an apical projection bearing 5 large spines
distally and a small inner process; distimere short and broad for most of
its length and with a pointed lateral projection at apex; aedeagus large, of
the uncus type; paraproct very long and tapered; a pair of elongate struc-
tures arising between the paraprocts (I cannot determine from what these
structures arise).
FEMALE, LARVA and PUPA. Unknown.
TYPE DATA. Holotype male, Sibulan River, 7000-8000 ft., Mt.
Apo, Davao, Mindanao, PHILIPPINES, in the Museum of Comparative
Zoology, Harvard. Paratype male with same date and locality as holotype.
DISTRIBUTION. Known only from the type locality.
TAXONOMIC DISCUSSION. A. nubicola can be mistaken for uncus
and was figured by Bohart (1945:fig. 50) as the latter species. The charac-
ters of the male terminalia, however, are distinctive as noted above and
shown in the figure.
BIOLOGY. Unknown.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 31
AEDES (NEOMACLEAYA) PAHANGI N. SP.
(Figure 23)
Species with wholly brown abdominal terga and sterna; it is ex-
ternally similar to hamistylus, robertsi or indecorabilis from which pahangi
can be differentiated by the shape of the preatrial plate and by the reticulate
spermathecae of the female as shown in the figure. The male, larva and
pupa are not known.
FEMALE. Head. Vertex dark with few pale scales around eye
margin and sides, a few upright scales confined to occiput; torus with some
hairs and flat scales; palpus about 1/6 length of proboscis; proboscis about
as long as fore femur, dark brown. Thorax. Scutal scales reddish brown;
anterior pronotum with 6 bristles and 2 short setae, without scales or these
possibly rubbed off; posterior pronotum with 4 bristles, 2-3 narrow curved
scales; a few pale scales on sternopleuron and upper mesepimeron; 6 bris-
tles on upper sternopleuron behind pale scales, without hairs on anterior
portion; spiracular area with 2-3 bristles, without scales; only a few mes-
epimeral hairs; lower mesepimeron bare; metameron bare; without scales
on propleuron. Wing. Alula fringed with broad dark scales. Legs. Fore-
coxal scales mostly rubbed off, the remaining ones pale brown; mid coxa
with a few pale scales; none on hind coxa; tarsal claws of all legs equal, each
claw toothed. Abdomen. Terga and sterna wholly brown with a few scattered
pale brown scales laterally. Terminalia. (Figure 23). Cercus long and
tapered, more than twice as long as basal width; postgenital plate longer
than wide, truncate distally; postatrial sclerite rounded, weak; postatrial
plate elongate ovate with a pair of sclerotized rods laterally; preatrial plate
pointed distally, hairy, a pair of platelets laterally; preatrial sclerite folded
over postatrial plate, with a deep median emargination. Three unequal sper-
mathecae, reticulate, without necks.
MALE, LARVA and PUPA. Unknown.
TYPE DATA. Holotype female, Kuala Terengan, Pahang, MALAYA
(J. L. Gressitt), 17. xii. 1958, in the Bishop Museum, terminalia and tarsal
claws on slides. Paratype 1 female (USNM), terminalia on slide, same date
and locality as holotype.
DISTRIBUTION. Known only from type locality.
TAXONOMIC DISCUSSION. This species is apparently closely re-
lated to hamistylus, robertsi or indecorabilis by the structures of the female
terminalia, by the toothed tarsal claws as well as other characters men-
tioned above.
BIOLOGY. Unknown.
32 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
AEDES (NEOMACLEAYA) PANAYENSIS LUDLOW
(Figures 4, 23)
Aedes panayensis Ludlow 1914, Psyche, Camb., Mass. 21:159 (o, 9);
Laffoon 1946, J. Wash. Acad. Sci. 36:242 («*, 9*); King and
Hoogstraal 1947, J. Wash. Acad. Sci. 37:119 (c).
A. panayensis differs from all other species in the subgenus by
having the fore and mid tarsal claws of both sexes equal, each claw toothed,
the hind claws equal, simple. On the basis of the male terminalia it appears
related to prioekanensis or dermajoensis as discussed under the latter spe-
cies; to incertus in the female. Laffoon (1946:242) and King and Hoogstraal
(1947:119) have given detailed descriptions of the species. The larva and
pupa are not known. |
FEMALE. Head. Vertex dark with a few pale scales on midline
and eye margin, pale lateral patches, sometimes narrow pale scales mixed
along midline; torus with few dark scales. Thorax. Scutal scales golden to
dark brown, lighter on front margin, above wing base and prescutellar area;
anterior pronotum with a few pale scales, 8 bristles; posterior pronotum
with narrow brown scales; patches of pale scales on upper and lower sterno-
pleuron, without hairs on anterior portion; hairs behind pale scale patch on
mesepimeron extend downwards to lower part of sclerite; metameron bare.
Legs. Fore and mid tarsal claws equal, each claw toothed; hind claws equal,
simple. Abdomen. Terga with pale lateral patches extending slightly on to
dorsum; sterna brown on apical half. Terminalia. (Figure 23). Cercus
gently tapered distally, about twice as long as basal width; postgenital plate
broad with a shallow emargination; postatrial sclerite flat at top; postatrial
plate small, pocket-shaped with wrinkled edges; preatrial plate consisting
of 2 small poorly sclerotized parts weakly connected medianly. Two large,
1 small spermathecae each with a small neck.
MALE. General habitus and legs as in female. Terminalia.
(Figure 23). Basimere with a narrow flattened apical projection serrated at
tip, 3 very strong setae close to inner dorsal margin, a long tapered pro-
jection arising from base; distimere curved distally, with 1-2 stout bristles
and weak setae beyond middle; paraproct broad, pointed distally.
LARVA and PUPA. Unknown.
TYPE DATA. Male lectotype selected by Laffoon (1946:242),
Iloilo, Panay, PHILIPPINES, in the U. S. National Museum.
DISTRIBUTION. Specimens examined: PHILIPPINES: Jolo, Jolo,
67 females, 18 males; Palawan, Puerto Princesa, 5 females; Leyte, 2 males,
2 females; NORTH BORNEO: Tarawakan, 1 female; NEW GUINEA: Schouten
Is., 26 females (USNM). Also recorded from the MOLUCCAS but I have not
seen specimens from this locality.
TAXONOMIC DISCUSSION. The female terminalia are remarkably
similar to those of incertus except that the edges of the pocket-shaped post-
atrial plate are wrinkled in panayensis, smooth in incertus. The male is
quite distinct as discussed above.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 33
BIOLOGY. Laffoon (1946:243) records that this species is ap-
parently confined to coastal areas. The adults have been collected biting
man during the daytime and resting in a crab hole in mangrove swamps.
The larvae have been taken in slightly brackish water in a beached canoe
and in a shaded, leafy, slightly brackish puddle in a nipa palm swamp at the
extreme upper limit of the tidal zone.
AEDES (NEOMACLEAYA) PHILIPPINENSIS N. SP.
(Figure 24)
A. philippinensis is a very distinct species quite unlike any other
species placed in Neomacleaya. The male basimere is short and simple, it
lacks apical or subapical projections but with a patch of strong spines and
setae on inner margin; the female postatrial plate has characteristic branch-
ed hairs and the preatrial plate consists of 2 widely separated, long, narrow
parts. The larva and pupa are not known.
FEMALE. Head. Scales of vertex all blackish brown, no visible
upright scales on occiput; torus with a few fine hairs and small flat scales;
palpus about 1/8 length of proboscis; proboscis longer than fore femur by
twice length of labella, dark brown. Thorax. Scutal scales uniformly dark
reddish brown; anterior pronotum with 4-5 strong bristles and a few short
setae, without scales; posterior pronotum with 4-5 bristles and a few scat-
tered narrow curved scales; postspiracular area with 2-3 bristles, without
scales; very few translucent scales on sternopleuron and upper mesepimeron;
4 bristles behind upper scales on sternopleuron, without hairs on anterior
portion; a few mesepimeral hairs behind translucent scales; lower mesepi-
meron bare; metameron bare; propleuron without scales, only bristles.
Wing. Alula with flat brown scales. Legs. Forecoxal scale patch mostly
brown with a small whitish spot above; mid coxa with some whitish scales;
no scales on hind coxa; fore and mid tarsal claws equal, each claw toothed;
hind claws equal, simple. Abdomen. Terga dark brown with small pale
lateral patches (these appear pale brown or translucent in holotype speci-
men); sterna wholly brown. Terminalia. (Figure 24). Cercus tapered
distally, about twice as long as basal width; postgenital plate rounded dis-
tally; postatrial sclerite produced medianly; postatrial plate with character-
istic branched hairs, wide opening and lateral projections; preatrial plate
consisting of 2 widely separated, long, narrow, more or less parallel-sided
parts. One large and 2 small spermathecae each with a fairly long neck.
MALE. The association of the sexes here is quite presumptive
due to lack of individual rearings. Although similar to female in general
habitus the male vertex has some pale scales laterally and a few upright
scales on occiput; torus bare; abdominal terga dark brown with pale brown
or translucent lateral patches. Legs. Fore and mid tarsal claws unequal,
only the larger claw toothed; hind claws equal, simple. Terminalia.
(Figure 24). Basimere short and simple, lacking apical and subapical pro-
jections but with a patch of strong spines and setae on inner margin; disti-
mere elbowed near base, narrowed distally with a short hook at apex; aedea-
gus well developed, of the andamanensis type; paraproct very short.
34 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
LARVA and PUPA. Unknown.
TYPE DATA. Holotype male, terminalia on slide, 10. viii. 1964,
Mahaplag, Leyte, PHILIPPINES (M. D. Delfinado), in the Bishop Museum.
Allotype female, terminalia and tarsal claws on slides, 25. ix. 1945,
Tacloban, Leyte (H. R. Roberts), in the U. S. National Museum.
DISTRIBUTION. PHILIPPINES.
BIOLOGY. A female has been bred out from a larva collected in
a rain pool; the male has been collected at light trap.
AEDES (NEOMACLEAYA) PRIOEKANENSIS BRUG
(Figure 8)
Aedes (Aedes) prioekanensis Brug 1931, Tijdschr. Ent. 74:249 (o*).
Species with wholly dark brown abdominal terga and toothed hind
tarsal claws; the male can be recognized by having a short blunt apical pro-
jection and 2 large spines on inner margin near base of basimere. The
female, larva and pupa are not known. The following description is based
on the type male in the British Museum. The specimen is in a very poor
condition.
MALE. Head. Lost. Thorax. Scutal scales brown; anterior
pronotum and propleuron destroyed; sternopleuron badly damaged by the
pin; posterior pronotum with 3 bristles, without scales; postspiracular area
with bristles missing; some pale scales present on upper mesepimeron; a
few hairs present behind this scale patch; lower mesepimeron and meta-
meron bare. Wing. Scales on alula missing. Legs. Few brown scales on
fore coxa; some pale scales on mid coxa; none on hind coxa; fore and mid
tarsal claws unequal, only the larger claw toothed; hind claws slightly un-
equal, each claw toothed. Abdomen. Terga wholly dark brown; sterna ap-
pear to have pale scales basally. Terminalia. (Figure 8). Based ona
slide mount from Sumatra. Basimere with a short blunt apical projection
and 2 large spines on inner margin near base; distimere twice bent, swollen
in basal 1/2, narrowed distally; aedeagus very simple as in dermajoensis ;
paraproct short and slender.
FEMALE, LARVA and PUPA. Unknown.
TYPE DATA. A male labeled type in the British Museum, ter-
minalia and tarsal claws in celluloid mount on pin, Benkulen, SUMATRA.
DISTRIBUTION. Known only from type locality.
TAXONOMIC DISCUSSION. On the basis of the male terminalia
prioekanensis appears to be closely related to dermajoensis and panayensis.
But the hind tarsal claws of prioekanensis differ from the 2 species men-
tioned in being slightly unequal, each claw toothed (Figures 8, 23); the
fore and mid claws unequal, only the larger claw toothed. In panayensis,
the fore and mid claws equal, each claw toothed; unequal,both large and
small claw toothed in dermajoensis; also prioekanensis has wholly dark
brown abdominal terga.
BIOLOGY. Brug (1931:249) records that the type male was bred
from a larva found in a swamp in a virgin forest.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 35
AEDES (NEOMACLEAYA) RARUS N. SP.
(Figure 24)
Although this species is based only on a unique male specimen
from North Borneo, the terminalia and tarsal claws are particularly dis-
tinctive. Like philippinensis, the basimere is short and lacks apical or
subapical projections and the aedeagus is of the andamanensis type. A.
varus, however, is distinguished by the following: Antenna strongly plumose;
anterior part of sternopleuron with fine hairs; lower mesepimeron bare;
metameron bare; fore and mid tarsal claws unequal, both large and small
claw toothed; hind claws simple, equal; abdominal terga with pale lateral
patches. Terminalia. (Figure 24). Basimere without apical projection
but with 2 large spines near base and a median flattened projection with
serrated tip; distimere expanded distally, with pointed apex; aedeagus of
the andamanensis type; paraproct very small, poorly developed.
FEMALE, LARVA and PUPA. Unknown.
TYPE DATA. Holotype male (British Museum), terminalia, mid
and hind claws on slide, B60/19, Tawau, NORTH BORNEO, ii. 1960 (D. H.
Colless).
DISTRIBUTION. Known only from type locality.
BIOLOGY. Unknown.
AEDES (NEOMACLEAYA) ROBERTSI LAFFOON
(Figures 13, 25)
Aedes (Aedes) robertsi Laffoon 1946, J. Wash. Acad. Sci. 36:230 (o*, 9*, L).
Adults of A. vobertsi are very similar to hamistylus from which
they can be separated by the details of the terminalia and male tarsal claws
as noted below. The larva, like hamistylus, has antennal hair 1-A single,
very long anal gills and a small slender siphon. The pupa is not known.
FEMALE. Head. Vertex dark brown with pale lateral patches,
2-3 pale scales between the eyes, dark upright scales on occiput; torus with
a few flat dark scales. Thorax. Scutal scales golden to dark brown in some
lights; anterior pronotum with some pale scales which may be rubbed off;
posterior pronotum with bristles, without scales; postspiracular area with
bristles, without scales; upper and lower sternopleuron with small patches
of loose pale scales, anterior portion without hairs; upper mesepimeron
without hairs behind pale scales; lower mesepimeron bare; metameron bare.
Legs. Tarsal claws of all legs equal, each claw toothed. Abdomen. Terga
wholly brown (occasionally a small pale or whitish lateral spot present on
tergum I); sterna entirely brown. Terminalia. (Figure 25). Cercus long,
tapered distally, more than twice as long as basal width; postgenital plate
longer than wide, with a shallow median emargination; postatrial sclerite
poorly sclerotized, flat at top; postatrial plate ovate with comparatively
large opening; preatrial plate large, rounded and open laterally, hairy;
preatrial sclerite folded over postatrial plate as a distinctive M-shaped
36 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
structure. One large and 2 small spermathecae each without a neck.
MALE. General habitus as in female. Legs, Fore and mid
tarsal claws unequal, only the larger claw toothed; hind claws equal,
toothed. Terminalia. (Figure 13). Basimeres joined subapically, rounded
apically bearing a small, curved, blunt process; distimere much swollen at
middle, abruptly narrowed and curved distally, bent at tip; aedeagus long
and tapered; paraproct rather stout throughout its length.
LARVA. (Figure 25). Extremely similar to hamistylus. Head.
1-C slender; 3-C very small; 4-C with 3-5 short branches; 5,6, 7-C with
2-3 branches; 8-C with 3 long branches; 9-C with 4-5 short branches; 10-C
with 2-3 branches, longer than 9-C; antenna spiculate; hair 1-A inserted at
about middle, single; 2-A longest of the apical hairs; mental plate with 32-
38 lateral teeth. Thorax. 1,2-P single; 3-P 3-4 branched; 8-P single; 9,
10,12-P 2-branched, barbed; 1-M single, long; 2-M very small, single; 3,
4-M with 3-5 long branches; 1-T very small, single; 2-T with 3 long
branches; 3-T with 4-5 branches; 4-T small branched tuft; 5-T very small,
single; 6-T with 2-3 long branches. Abdomen. 6-I-VI single; 7-I single;
7-II-VI with 3-4 branches; segment VIII with 9-12 comb scales, each scale
pointed and with a fine lateral fringe; 1, 4-VIII single; 2-VIII double; 3, 5-VIII
with 3-4 barbed branches; siphon small and slender, about 3 times as long
as basal width, with 5-8 pecten teeth, spacing between the teeth nearly
regular, last tooth inserted about 2/3 from base; siphonal tuft 1-S with 1-3
branches; inserted near apex; saddle fairly smooth; 1-X-1-2 branched; 2-X
with 2-3 branches; 3-X single; anal gills very long, about 6 times as long as
saddle; 1-2 precratal tufts.
PUPA. Unknown.
TYPE DATA. Holotype male, Tacloban, Leyte, PHILIPPINES,
in the Academy of Natural Sciences, Philadelphia. Paratypes 8 males, 11
females, 6 larvae, with same data as holotype in the U. S. National Museum
and British Museum (Natural History).
7 DISTRIBUTION. Specimens examined: PHILIPPINES: Leyte,
Tacloban, 6 males, 8 females (USNM), 14 larvae (ANS).
BIOLOGY. Larvae were collected in gravel-bottomed residual
pools ina stream by H. R. Roberts. Nothing is known of the biting habits.
AEDES (NEOMACLEAYA) SIAMENSIS N. SP.
(Figure 26)
This species is very distinct externally by the presence of numer-
ous broad white scales on anterior pronotum, postspiracular area and in
front of vertex extending downwards to frons. The male basimeres are
joined subapically, rather short with small inner and outer apical projec- —
tions; the female preatrial and postatrial plates resemble those of pahangi.
The larva has antennal hair 1-A single and large siphon; the pupa has gener-
ally weak hairs but 2-I-VI heavy, almost spine-like.
FEMALE. Head. Front of vertex with a patch of broad white
scales extending downwards to frons, a few pale scales laterally and on
Delfinado: Aedes (Neomacleaya) in Southeast Asia 37
occiput; torus with some small dark scales on inner side; palpus about 1/8
length of proboscis; proboscis about as long as fore femur, dark. Thorax.
Scutal scales blackish; anterior pronotum covered with broad white scales,
0-6 bristles; posterior pronotum with 5-6 bristles posteriorly, 3-4 loose
pale scales; postspiracular area covered with white broad scales, 3-4 short
bristles; large patches of white scales on upper and lower sternopleuron
and upper mesepimeron, anterior portion of sternopleuron without hairs;
lower and posterior portion of mesepimeron bare; metameron bare; pro-
pleuron covered with white scales. Wing. Alula fringed with lanceolate
scales. Legs. Forecoxal scale patch all white scaled; mid and hind coxae
each with small patches of white scales; femora with extensive pale scaling
on ventral and basal posterior surfaces; tarsal claws of all legs equal, each
claw toothed. Abdomen. Terga usually with pale lateral patches produced
on to dorsum but not forming complete bands; sterna pale scaled on basal
1/2. Terminalia. (Figure 26). Cercus long and slender, about 3 times as
long as basal width; postgenital plate longer than wide, truncate or slightly
emarginate distally; postatrial sclerite very weak (not shown in the figure);
postatrial plate with a large opening and a brace-like process on top; pre-
atrial plate similar to pahangi, pointed distally, hairy, but lacks lateral
platelets; preatrial sclerite shown overlying the postatrial plate in the figure.
Two large, 1 small spermathecae with very short, almost indistinct necks.
MALE. Externally similar to female except the abdominal terga
which are wholly brown though sometimes with some pale brown or whitish
scales scattered laterally; sterna mostly pale; posterior basal surface of
femora dark. Legs. Fore and mid tarsal claws slightly unequal, the larger
claw with a double tooth; hind claws toothed, subequal; this is apparently
variable as shown in figure 26. Terminalia. (Figure 26). Basimeres
joined subapically, rather short, with a short outer apical and 2 small inner
projections; distimere expanded and hairy at middle, abruptly narrowed and
curved distally, bent at tip; aedeagus simple, pointed apically; paraproct
slender, tapered and pointed distally.
LARVA. Figure 26 and description based on cast skins of allo-
type and paratypes. Head. 1-C slender; 3-C very small; 4-C with 3-5 weak
short branches which are difficult to see; 5,6-C with 1-3 branches; 7-C with
3-5 branches; 8-C with 1-2 branches; 9,10-C with 3-5 branches; 12,14-C
Single, very small; mental plate with 32-34 lateral teeth; antenna spinose;
1-A inserted before middle, single; 2-A longest of the apical hairs. Thorax.
1,3-P single; 2-P single or double; 8-P single; 9,10-P single; 12-P double;
1-M with 2 fairly long branches; 2-M short, single; 3, 4-M with 3-4 long
branches; 1-T single; 2,3-T with 5-6 long branches; 5-T very small, single;
6-T long, single. Abdomen. 6-I double; 6-II-VI single; 7-I single; 7-II-VI
with 3 or more short branches; segment VII with 7-10 comb scales, each
scale pointed and with lateral fringe; 1-VIII with 3-4 short branches; 2-VII
double; 3- VIII with 5-8 barbed branches; 4-VIII single; 5-VIII with 4-6
barbed branches; siphon large and stout, about twice as long as basal width,
rugose, with 12-14 pecten teeth, each tooth with a basal denticle except last
tooth which is usually simple; siphonal tuft 1-S with 2-5 branches, inserted
2/3 from base; saddle broader than long, distal margin smooth; 1-X single;
38 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
2-X with 3-4 branches; 3-X single; anal gills more than twice length of
saddle; 2-3 precratal tufts.
PUPA. Body hairs generally weak and short. Cephalothorax.
1,2,3-C 2-branched; 10-C with 3-4 branches; 11-C single; 12-C with 2-3
branches distally. Abdomen. 2-I-VI fairly heavy, almost spine-like; 3-I
with 3-4 branches; 3-II-III single; 1, 3,5-IV-VII at most 2-branched; 6-I
double; 6-II single, as long as 6-I; 6-ID-VII single, short; 9-VII single,
fairly long; 9-VIII single; 10-III-VII single, long; 7-VI-VII single, long;
T-III-V branched. Paddle. Outer margin serrated, with strong midrib;
1-P simple. :
TYPE DATA. Holotype male with associated pupal skin (01998-
103), allotype female with associated larval and pupal skins (01998-3),
terminalia and tarsal claws on slides, Lampang, Ban Pong Tao, THAILAND,
21.iv.1965; paratypes 11 males, 14 females with 11 pupal and 2 larval skin
mounts, with same date and locality as holotype. Male holotype, female
allotype, 6 male, 6 female paratypes in the U. S. National Museum; 3 male,
4 female paratypes in the British Museum (Natural History); 2 male, 4
female paratypes with associated skins in the Bishop Museum.
DISTRIBUTION. Known only from type locality.
TAXONOMIC DISCUSSION. There is a marked sexual dimorphism
in stamensis. Individual rearings of adults from gravid females caught at
Ban Pong Tao definitely associate the sexes through the immature stages.
The development of teeth on the larger tarsal claw of the male fore and mid
legs seems to be extremely variable.
BIOLOGY. Possibly a ground pool breeder; the females were
collected biting man.
AEDES (NEOMACLEAYA) SINGULARIS (LEICESTER)
(Figure 22)
Aioretomyia singularis Leicester 1908, Cul. Malaya 3:188 (c’).
Aedes (Aedes) singularis (Leicester), Edwards 1917, Bull. ent. Res. 7:222,
fig. 8c (o'*); Edwards 1922, Indian J. med. Res. 10:265 (c, key).
A dark reddish brown species with numerous hairs on lower mes-
epimeron and pale lateral patches on abdominal terga. It is particularly
distinguished by the many strong bristles behind pale scale patch on sterno-
pleuron which extend downwards to lower portion; by the bifurcate distimere
of the male terminalia and by the unusual shape of the postatrial and pre-
atrial plates of the female terminalia. The larva and pupa are not known.
FEMALE. Head. Vertex blackish brown with patches of pale
scales laterally, numerous upright scales on occiput and some narrow
curved golden ones at middle continuing to occiput; torus with a few small
dark scales; palpus 1/6 - 1/8 length of proboscis; proboscis as long as or
slightly longer than fore femur, dark brown. Thorax. Scutal scales red-
dish brown, golden along front margin; anterior pronotum with 6 bristles on
top, 4 short ones below, without scales; posterior pronotum with 4 bristles,
Delfinado: Aedes (Neomacleaya) in Southeast Asia 39
some narrow curved scales; postspiracular area with 3-4 bristles, without
scales; small patches of pale scales on upper and lower sternopleuron, with
many strong bristles and short hairs behind upper scale patch which extend
to lower portion; upper mesepimeron with numerous fine hairs behind pale
scale patch which extend downwards to lower portion of sclerite; metameron
almost bare, with some fine hairs very difficult to see; propleuron largely
brown with a small pale spot above. Wing. Alula fringed with narrow dark
scales. Legs. Forecoxal scale patch mostly brown; a row of pale scales
each on mid and hind coxae; fore and mid tarsal claws equal, each claw
toothed; hind claws simple, equal. Abdomen. Terga with large pale lateral
patches, those on VI-VII with ends produced on to dorsum; sterna pale
scaled on basal 2/3, brown apically. Terminalia. (Figure 22). Cercus
tapered distally, more than twice as long as basal width; postgenital plate
small, with a fairly deep emargination; postatrial sclerite indistinct; post-
atrial plate well sclerotized, with a large uncus-type opening and long hairs;
preatrial plate entire, broader distally, hairy. One large and 2 small sub-
equal spermathecae without necks.
MALE. Similar to female in general habitus. Legs. Fore and
mid tarsal claws unequal , only the larger claw toothed; hind claws equal,
simple. Terminalia. (Figure 22). Basimere produced sternally with an
apical projection dentate at tip and a small basal process; distimere bifur-
cate at distal half, the shorter branch with a small lateral tooth; aedeagus
large, of the uncus type; paraproct long and slender, evenly tapered, curved
inwards. Sternum IX projecting distally, with a deep median concavity.
LARVA and PUPA. Unknown.
TYPE DATA. A male labeled type in the British Museum, termi-
nalia supposedly in celluloid mount on pin, but missing, Kuala Lumpur,
MALAYA,
DISTRIBUTION. Specimens examined: MALAYA: Kuala Lumpur,
1 male (type); SARAWAK: Kuching, 3 females, 2 males (BM). :
TAXONOMIC DISCUSSION. This species superficially resembles
virilis but the structure of the aedeagus and the postatrial plate of the female
resemble those of uncus; singularis, however, is a very distinct species and
can be readily separated by the characters mentioned in the key.
BIOLOGY. Unknown.
AEDES (NEOMACLEAYA) VARIETAS (LEICESTER)
(Figure 10) |
Aioretomyia varietas Leicester 1908, Cul. Malaya 3:185 (¢ only).
Aedes (Aedes) varietas (Leicester), Edwards 1917, Bull. ent. Res. 7:222,
fig. 8h (¢*); Knight and Hull 1953, Pacif. Sci. 7:474 (type 2 =
uncus).
Species with pale lateral patches on the abdomen, bare lower
mesepimeron and simple hind tarsal claws. The male is readily distin-
guished from that of other species by the characteristic shape of the
40 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
distimere as shown in the figure; the basimere has forked apical and sub-
apical projections and 3 long curved spines at the base. The female is as-
sociated here with uncertainty but I have the terminalia illustrated as in
figure 10; the larva and pupa are not known.
MALE. Head. Vertex dark brown with pale scales laterally, a
few upright scales mixed with some pale ones on occiput; torus bare; palpus
about 1/10 length of proboscis; proboscis about as long as fore femur, dark
brown. Thorax. Scutal scales mostly reddish brown; anterior pronotum
with 4 bristles, without scales; posterior pronotum with 3-4 bristles; post-
spiracular area with 3 bristles, without scales; upper and lower sterno-
pleuron with small patches of pale scales, without hairs on anterior portion,
5 bristles behind upper scale patch; small patch of pale scales on upper
mesepimeron, only a few hairs behind this scale patch; lower mesepimeron
bare; metameron bare; propleuron with pale scales. Wing. Alula fringed
with narrow dark scales. Legs. Forecoxal scale patch mostly brown; mid
and hind coxae each with some pale scales; fore and mid tarsal claws un-
equal, only the larger claw toothed; hind claws small, equal, simple.
Abdomen. Terga with patches of pale scales laterally; sterna light brown
scaled. Terminalia. (Figure 10). Basimere with a divided inner subapical
projection, a forked apical one with 3-4 prongs, 3 long slender curved
spines at base; distimere distinctive by being roughly wedge-shaped in
which one point is narrowly prolonged and bent; aedeagus fairly large, of
uncus type; paraproct long and slender, evenly tapered.
FEMALE. Uncertain; 3 specimens from Malaya labeled varieias
in the British Museum are provisionally placed here. It agrees well with
the male except: anterior pronotum with 6-8 weak bristles; 4-6 bristles on
postspiracular area; fore and mid tarsal claws equal, each claw toothed;
hind claws equal, simple. Terminalia. (Figure 10). Cercus tapered
distally, about twice as long as basal width; postgenital plate broader than
long, with a fairly deep median emargination; postatrial sclerite with lobed
lateral corners, convex medianly; postatrial plate with a large opening and
long fine hairs; preatrial plate large, subquadrate, weakly divided, hairy.
Two large and one small spermathecae each with a short neck.
LARVA and PUPA. Unknown.
TYPE DATA. A male; in the British Museum, labeled "cotype/
Jungle Pahang Rd 5th mile/midday/Kuala Lumpur/7/11/03"; terminalia
supposedly in celluloid mount on pin, but missing. Since this is the only
male I assume it is the male lectotype selected by Knight and Hull (1953:
474) but it is not so labeled ; the females were assigned to uncus.
DISTRIBUTION. Specimens examined: SUMATRA: Kota Tjane,
1 male; MALAYA: Kuala Lumpur, Klang Jungle, 3 females (BM).
TAXONOMIC DISCUSSION. The association of the sexes here is
only presumptive; it is likely that they are correctly associated since the
general anatomy and coloration are similar. The structures of the male
terminalia superficially resemble those of cautus and atrius by having long,
curved inner spines on the basimeres; the large preatrial plate of the
female resemble atrius, the postatrial plate and sclerite is like cautus.
varietas is easily differentiated from both by the shape of the distimere and
Delfinado: Aedes (Neomacleaya) in Southeast Asia 41
by the forked projections of the basimere in the male, as well as by the
details of the postatrial and preatrial plates in the female.
BIOLOGY. Unknown.
AEDES (NEOMACLEAYA) VIRILIS (LEICESTER)
(Figure 22)
Verrvallina virilis Leicester 1908, Cul. Malaya 3:197 (c).
Aedes (Aedes) virilis (Leicester), Edwards 1917, Bull. ent. Res. 7:222,
fig. 8b (o'*).
There is no specimen available for study except the type in the
British Museum. The specimen is in very poor condition; the head is mis-
sing, the abdomen is broken off and glued to a card and there are no tarsal
claws; its sex cannot be determined. On another pin bearing this name and
labeled type is a mount on celluloid of a male terminalia similar to those
figured by Edwards (1917:222, fig. 8b). I assume that these terminalia
came from the type; the external characters, such as still remain, agree
well with Leicester's description. )
MALE. Thorax. Scutal scales brown to reddish brown; post-
spiracular area with 2 bristles; sternopleuron with a few hairs on anterior
portion, 7 bristles behind upper scale patch, a small patch of scales on
lower portion mixed with some hairs; upper mesepimeron with numerous
hairs behind pale scale patch which extend downwards to lower mesepimeron;
metameron bare; propleuron with some pale scales. Wing. Remaining
fringed scales on alula narrow. Legs. Forecoxal scale patch mostly brown
with a few pale ones scattered below; mid and hind coxae appear bare;
Leicester (1908) describes "fore and mid ungues unequal the larger ungues
uniserrate", but did not mention the hind claws. Abdomen. Terga with pale
lateral patches; sternal scales mostly rubbed off with a few pale ones left.
Terminalia. (Figure 22; after Edwards 1917). Due to its poor preparation
it has not been possible to redraw the type specimen. Basimere with a short
slender apical projection, a shorter inner projection divided at tip, a small
process and a hairy lobe at base which are not evident in Edward's figure;
distimere particularly distinctive by being very long, tapered distally, with
a small pointed process at middle; paraproct long and slender; aedeagus
difficult to follow but appears similar to singularis.
FEMALE, LARVA and PUPA. Unknown.
TYPE DATA. Type male (terminalia) in the British Museum,
with the following locality data: 'Jungle 5th mile/Gombak Rd/Kuala Lumpur/
FED. MALAY STATES."
DISTRIBUTION. Known only from type locality.
BIOLOGY. Unknown.
42 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
SPECIES INQUIRENDA
Atoretomyia perdita Leicester 1908, Cul. Malaya 3:192 (o’). Gombak Road,
Kuala Lumpur.
The identity of this species is doubtful. The type has been lost
and the description of Leicester was made up from his recollection. The
presence of 2 long sharp spines at the base of larger fore tarsal claws as
well as a number of characters he mentioned certainly do not fit in this
subgenus.
CORRECTIONS TO AEDES (NEOMACLEAYA) IN THAILAND
Page 11, line 48: Change the larger claw toothed to both claws toothed as in
figure 24.
Figure 12, A. incertus, larva. ee hair ie 9-P should be
12-P
ACKNOWLEDGEMENTS
As in the past my sincere thanks are again due to Dr. Alan Stone
and Dr. Botha de Meillon for their many helpful suggestions regarding this
revision and for reviewing the manuscript. I am also particularly indebted
to Dr. P. F. Mattingly and Dr. J. Bonne-Wepster for much kindness to me
in London and Amsterdam and for their helpful information concerning the
Oriental species. The illustrations were drawn by Mrs. Elaine R. Hodges
and Mrs. Jung Lea Smith of Southeast Asia Mosquito Project and by the
artists of the 406th Medical Laboratory, Tokyo. I am especially apprecia-
tive of their assistance. Miss Helle Starcke typed the manuscript.
LITERATURE CITED
BARRAUD, P. J.
1928. A revision of the culicine mosquitoes of India. Part XXIV -
The Indian species of the subgenera Skusea and Aedes, with
descriptions of eight new species, and remarks on a new
method for identifying the females of the subgenus Aedes.
Indian J. med. Res. 16: 357-375, illus.
1934. Family Culicidae. Tribes Megarhinini and Culicini. The
Fauna of British India, including Ceylon and Burma.
Diptera. Vol. V, 463 pp. , illus. London.
Delfinado: Aedes (Neomacleaya) in Southeast Asia 43
BOHART, R. M.
1945. A synopsis of the Philippine mosquitoes. U. S. Navmed 580,
88 pp., illus. Washington, D. C.
1956. New species of mosquitoes from the southern Ryukyu Islands.
Bull. Brooklyn ent. Soc. 51: 29-34, illus.
BRUG, S. L.
1931. New Culicidae from Sumatra. Tijdschr. Ent. 74: 245-250,
illus.
DYAR, H. G. and R. C. SHANNON.
| 1925. The types of Philippine mosquitoes described by Ludlow and
other notes on the fauna (Diptera, Culicidae). Insec. Inscit.
menst. 13: 66-89.
DELFINADO, M. D.
1967. Contributions to the mosquito fauna of Southeast Asia. I. The
genus Aedes, subgenus Neomacileaya Theobald in Thailand.
Contr. Amer. ent. Inst. 1(8): 1-35, illus.
EDWARDS, F. W.
1917. Notes on Culicidae, with descriptions of new species. Bull.
ent. Res. 17: 201-229, illus.
1922. A synopsis of adult Oriental culicine (including megarhinine
and sabethine) mosquitoes. PartI. Indian J. med. Res. 10:
249-293.
KING, W. V. and H. HOOGSTRAAL.
1947. New Guinea species of mosquitoes of the genus Aedes, sub-
genus Aedes. J. Wash. Acad. Sci. 37: 113-134, illus.
KNIGHT, K. L. and W. B. HULL.
1953. The Aedes mosquitoes of the Philippine Islands Il. Subgenera
Aedemorphus, Aedes and Cancraedes (Diptera, Culicidae).
Pacif. Sci. 7: 453-481, illus.
LAFFOON, J.
1946. The Philippine mosquitoes of the genus Aedes, subgenus
Aedes. J. Wash. Acad. Sci. 36: 228-245, illus.
LEICESTER, G. F.
1908. The Culicidae of Malaya. Stud. Inst. med. Res. F. M. S.
3: 18-261.
LUDLOW, C. S.
1908. Mosquito notes, no. 6. Canad. Ent. 40: 50-52.
THEOBALD, F. V.
1910. Second report on the collection of Culicidae in the Indian
Museum, Calcutta, with descriptions of new genera and
species. Rec. Indian Mus. 4: 1-33, illus.
44 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
APPENDIX
PRESENT STATUS OF THE AEDES (NEOMACLEAYA) FAUNA
SPECIES
andamanensis Edwards
atrius Barraud
campylostylus Laffoon
cautus Barraud
clavatus Barraud
cretatus Delfinado
cyrtolabis Edwards
dermajoensis Brug
dux Dyar and Shannon
fragilis (Leicester)
gibbosus Delfinado
hamistylus Laffoon
hispidus Delfinado
incertus Edwards
indicus (Theobald)
indecorabilis (Leicester)
ly
unknown.
uncertain.
1 +
J
?
L
Pp
stage or sex described and illustrated.
stage or sex described but not illustrated.
OF SOUTHEAST ASIA 1
BIOLOGY
Larval habitats known
Larval habitats known
Larval habitats known
Adult biology and host prefer-
ence known
Unknown
Larval habitats known
Larval habitats known
Habitats known
Well known
Unknown
Larval habitats known
Larval habitats known
Unknown
Larval habitats known
Larval habitats known
Larval habitats known
Delfinado: Aedes (Neomacleaya) in Southeast Asia 45
SPECIES og Lee BIOLOGY
ishigakiensis Bohart * * * * Larval habitats known
johnsoni Laffoon * * * * Larval habitats known
latipennis Delfinado * * * * Larval habitats known
leicesteri Edwards * - - - Unknown
lugubris Barraud * * = - Unknown
macrodixoa Dyar and Shannon * * * - ~~ Larval habitats known
margarsen Dyar and Shannon * * * * \JLarval habitats known
neomacrodixoa King and
Hoogstraal * * = - Unknown
nigrotarsis (Ludlow) * * * * Larval habitats known
notabilis Delfinado - * - - Unknown
nubicola Laffoon * - - - Unknown
pahangi n. sp. - * = - Unknown
panayensis Ludlow * * = - Well known
philippinensis n. sp. ee Larval habitats known
prioekanensis Brug * - = = Larval habitats known
protuberans Delfinado * = - - Unknown
pseudodiurnus (Theobald) * - - = Unknown
vavus 1. sp. * - - - Unknown
robertsi Laffoon * * * - Larval habitats known
Siamensis Nn. Sp. * %* * + Host preference known;
larval habitats unknown
* * - - Unknown
singularis (Leicester)
46 Contrib. Amer.
SPECIES
torosus Delfinado
uncus (Theobald)
vallistris Barraud
varietas (Leicester)
virilis (Leicester)
Ent.
Inst.
, vol. 2, no. 4, 1968
P BIOLOGY
- Unknown
* Host preference and larval
habitats known
o Larval habitats known
- Unknown
=~ Unknown
_metameron
ANDAMANENS!/S
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>
KR
Nes
Ly
&
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onm ———4
Om
DUX
acai
mi
TYOSHIGAK)
LYOSHIGAKI
PANAVENS/S
1.0mm 1.0mm
UNCUS
/
paraproct
CAMPYLOSTYLUS
24 90% o,
4 Z
CAMPYLOSTYLUS
PRIOEKANENS/S
paraproct
FRAG/IL/S
PRIOEKANENSIS
paraproct
FRAG/L/IS
LEICESTERI
a =
‘ yy
AVG XR,
AV ta aes
‘i Vert OS yeti
end ;
OF
IR
oe
a TS 5 3,
HAMISTYLUS he
oa
paraproct
(itz
a
Noe
VARIETAS
WA
paraproct
hodges HAMISTYLUS
\
UMN yey i Fal ae
ISHIGAKIENS/S
Fig. 12
MP
1.0mm
=
SS
——
—
——
SZ
i. Sa
vill
\
ah’
MITA
0 Fe
\Yy <a = fies
iow
ISHIGAKIENS/S
hind
ela 0.05mm
e
O.lImm
paraproct
mid
claw
LUGUBRIS
Fig. 14
MACRODIXOA
extension of
IX tergum
NEOMACRODIXOA
MACRODI/XOA NEOMACROD/XOA
ag ge A ee a Na a Ste
MARGARSEN
ae
UV
1°)
oe
Qo.
12)
=
2}
a
MARGARSEN
0.05mm
1.0mm
paraproct
JOHNSON!
JL. SMITH
JOHNSONI/
ya - so ied dn me ree = in pA raph A ae My, ee ae
paraproct
N/IGROTARS/S
See aera _- ms 3 =
OT J+ Oh 0 fiend be NS ©
fs 5
. iy Mt
8 ,
—f
sso f& oe x
kiss
‘ ¢
(
\
\
y y |
j
/,
V1
LV
Mag
Hh
\
y
ii
NIGROTARSI/S
0.2mm
paraproct
TS) ae
Ss Uhl bo
PA Re
AN
TA AN
UU
v— rs =
SINGULARIS
paraproct
paraproct
(after Edw.
117: 222,
fig.8b)
NUBICOLA
claine
aa VIRILIS
Ve
Se}
0.05mm
DERMAJOENS/S
“A
PANAYENS/S ‘
i
SSR eee
Te SSS Ei ee
fa aa a
er ay WKS
4 iawiatnn)
2 \) an My
it
PANAYENS/S PAHANG/
elaine
r
hodges
“. paraproct
PH/ILIPPINENS/S
larva
Paratype
Ma
Qe
Wy
a
S)
ne
3& mid claw
paratype holotype
o' fore claw
preatrial
sclerite
S/IAMENSIS
elaine
hodges
Delfinado: Aedes (Neomacleaya) in Southeast Asia
|
Valid names are set in roman type, synonyms are italicized. The
italicized pages are those which begin the primary treatment of the species.
Numbers in parentheses refer to the figures illustrating the species in ques-
The first column refers to species mentioned in "Contributions to the
Mosquito Fauna of Southeast Asia - I.
Neomacleaya Theobald in Thailand" and the second column to ''Contributions
to the Mosquito Fauna of Southeast Asia - III. The Genus Aedes, Subgenus
tion.
INDEX
Neomacleaya Theobald in Southeast Asia. "'
Aedes
adustus
Aioretomyia
andamanensis
atrius
butleri
campylostylus
cautus
cinereus
clavatus
cretatus
Culex
_cyrtolabis
dermajoensis
dux
fragilis
gibbosus
hamistylus
hirsutipleura
hispidus
imitator
incertus
indecorabilis
indecorabilis
indica
indica var. simplex
indicus
ishigakiensis
johnsoni
latipennis
» 2)
2s 6, 1. 8, y, 10, 12,
9 a, 6, pe 8, pe 12, (3, 4)
ia
o
32
Mi CO OD
, 6, 8, 12, (18, 20)
,7,8, 16, (7)
or? @)
8,19, 20, (9, 10)
,5, 6, 7, 8, 20, (11, 20)
4,7, 8, 21, (18)
25,26
4, %,%, 8,22, 24, 12, 19)
25
4,5, 7, 8, 25, 26, (13, 20)
2,24,25
9
4,7, 8, 22, 24, 25, (18, 20)
10
4,5, 6, 8, 26, (14, 15)
The Genus Aedes, Subgenus
1
2,42
2,5,10,12, 14,19, 20, 25,
26, 28, 33, 35, (1)
2,6, 11, 12, 40, (24)
6, (9, 10)
3,6,12, 32, 42
3.9520, 16; 18, 38
Oouwp on
mee COD
,12, 13, 14, 25, (6, 7)
0
148, 28, 19) (11, 12)
0,12, 19, 25, 26,
714 Contrib. Amer. Ent. Inst., vol. 2, no. 4, 1968
leicesteri
lugubris
macrodixoa
macrodixoa
malayt
margarsen
medtofasciata
neomacrodixoa
nigrotarsis
notabilis
nubicola
pahangi
panayensis
perdita
philippinensis
-prioekanensis
protuberans
pseudodiurna
pseudodiurnus
rarus
robertsi
siamensis
sigmoides
singularis
Skusea
taentata
torosus
uncus
vallistris
varietas
varietas
Verrallina
virilis
yusafi
3,18
10
24,25
3
24,25
4,7, 8, 28, (18)
4,6, 8, 29, (19)
29
4,5, 6, 8, 29, (19)
18, 20
21
24,25, 29
22,23
4,5, 6,8, 30, (19)
4,5, 6,8, 30, 32, (16, 19)
4,5, 6, 8, 32, (17)
2,30
1,2, 24,25
2
Til
3, 9, 21, (8)
3, 4, 8, 22, (13)
3,5, 8, 12,23,24,277(14,
15)
2,15, 16
3,4, 5,10, 12,19, 20, 25, 26,
(16, 17)
, 24, 27, (15)
2, 28,29, (20, 21)
= ©
>
~
4,27, 30, (22)
1, 36, 37, (23)
, 8, 32, 34, (4, 23)
O1% DN
~
CW WDND RO & OW Ww
~~
3,5, 9, 38, 39, 41, (22)
3.9.44
3,5, 9, 21, 24, 27, 30, 39, 40,
(5)
3,6,11
3,6, 11, 39, 40, (10)
1
3, 9,39, 41, (22)
Be
ei
a Rae
Peet
Bisby
hogl
ely ae
oF
ELS
Ses
pea
pie
he
Eva
Contributions
of the
American Entomological Institute
Volume 2, Number 5, 1968
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA. - IV
Species. of the subgroup Chrysolineatus of group D,
genus Aedes, subgenus Finlaya Theobald
by
Kenneth L. Knight
CONTRIBUTIONS
of the
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Thailand.
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No. 5. Contributions to the mosquito fauna of southeast Asia.
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postpaid
Contributions
of the
American Entomological Institute
Volume 2, Number 5, 1968
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF
SOUTHEAST ASIA. - IV
Species of the subgroup Chrysolineatus of group D,
genus Aedes, subgenus Finlaya Theobald
by
Kenneth L. Knight
CONTENTS
Tt CU I a ears Ge ane GRE AIM MMe a be 1
CPOUp Wee URES ITAA TUS ea 8 RW ee Se ee 2
BuberTOouD L CMR YOOLINMAT US 6c ek ee ee ke ae 2
KEYS TO THE SPECIES
OO Ue PON PRA ek ee eS al ae 5)
PUTT ee, UE VEG poi ce ah ee oe er ie ee a 6
DESCRIPTIONS OF THE SPECIES
euyysaijeeaius (Theobald) . 6s. ee ee ee 8
PORT OC CHRIS PA ie a ee ek ee er we 10
Povey) (SALTON Fe ee ea 13
yaponirus joponious (Theobald) ..< 6. ek. ek ee 4 15
japonicus shintienensis Tsaiand Lien .......... 17
Tees (Cea) i we ee 19
RC ie pha ek WW ae ele howe 20
VR A a ee ee a bas oe
SOPOT OWA a ke wae ae ee ee we 29
Bee Oe a gk Be be a elec ke eared 27
PAR EN Pee es ele a eee ce ben ee ke 29
Ree CEE a Pa os ee ae eee a ee 29
a ee ee Re ee ke HU 1.12
Re ie ee ere oe re a ere el aoe 45
CONTRIBUTIONS TO THE MOSQUITO FAUNA OF SOUTHEAST ASIA. - IV.
SPECIES OF THE SUBGROUP CHRYSOLINEATUS OF GROUP D,
GENUS AEDES, SUBGENUS FINLAYA THEOBALD!
By
Kenneth L. Knight?
INTRODUCTION
This is the first of a series of revisionary papers on the Aedes
(Finlaya) mosquitoes of Southeast Asia. Groupings of species, believed to
be natural because of the common possession of numerous structural and
biological characters, will be sought within the subgenus and treated one by
one. The inter-relationships of these groupings will be described and dis-
cussed in the terminal paper. Until the appearance of that paper, the intra-
subgeneric classification of Knight and Marks (1952) will be used where pos-
sible.
Species which occur near to, but outside of, Southeast Asia are in-
cluded in the keys wherever possible. The larval and pupal chaetotaxy used
is that of Belkin (1962). An asterisk following the abbreviations used (2 =
female, « = male, L = larva, P = pupa) indicates that at least some portion
of that sex or stage is figured. All types of included species which are in
the British Museum (Natural History) and the United States National Museum
have been seen by me; Attention is drawn to the fact that the thickness of
many of the larger larval hairs have been over-exaggerated in figures 5, 6,
7, 8, 9 and 10.
Abbreviations used in the references to literature conform to the
World List of Scientific Periodicals, 3rd ed. , Academic Press, 1952.
, This work was supported by Research Contract No. DA-49-193-MD-2672
from the U. S. Army Medical Research and Development Command,
Office of the Surgeon General.
Department of Entomology, University of Georgia, Athens, Georgia.
30601.
2 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
GENUS AEDES MEIGEN
SUBGENUS FINLAYA THEOBALD
GROUP D, AUREOSTRIATUS
SUBGROUP I, CHRYSOLINEATUS
The classification used here is that proposed by Knight and Marks
(1952) who divided Group D into 8 subgroups of which 3 occur in Southeast Asia,
namely, Subgroup I, CHRYSOLINEATUS; Subgroup Il, AUREOSTRIATUS;
and Subgroup VI, TOGOL,
Group D, AUREOSTRIATUS
Tentatively distinguished by the following: scutal pale scaling in-
cludes a pattern of narrow longitudinal lines, this pattern in a few species
occasionally obliterated by a general diffusion of pale scales. Femora and
tibiae not lined anteriorly with pale scales for nearly their whole length.
Tarsi with basal pale scaling on some segments. Wings not extensively
spotted with areas of pale and dark scaling. Larval, pupal, and biological
characteristics will be added to this definition in subsequent papers.
Subgroup I, CHRYSOLINEATUS
The members of this subgroup have many characters in common.
Consequently, in order to simplify the species descriptions which follow,
these characters are summarized here and will not be repeated unless sub-
ject to extensive variation or used as species-separation characters.
FEMALE. Head. Torus with a patch of small broadened scales
mesally; palpus about one fifth to one fourth length of proboscis, dark scaled,
usually some pale scales apically; proboscis dark scaled, some species with
areas of pale scaling; vertex with a longitudinal median band of narrow pale
scales, this band widened posteriorly into a broad sparse patch, a line of sim-
ilar scales along median half of eye margins, a submedian area of dark
scales between ocular pale line and posterior patch of narrow pale scales;
lateral portion of vertex with broad pale scales, small anterior patch of dark
scales usually visible medially in this area; forked upright scales abundant
dorsally from eye margin to occiput and usually all dark. Thorax. Anterior
pronotum with a dense cover of broad pale scales, sometimes a few narrow
scales present; posterior pronotum variably scaled; propleuron with a patch
of broad pale scales, this patch continuing onto the anterior surface; scutum
with brown integument, covered with small narrow brown and pale (yellow
or golden) scales, pale scales arranged in narrow longitudinal lines as fol-
lows: a full length median line, a full length submedian line interrupted at
scutal angle, anterior end of posterior portion of this line usually curved
outward to a variable extent along scutal angle, and a line of variable extent
over wing base which mav be just a diffused area in some species, a small
patch of long narrow-curved pale scales just before wing base, narrow pale
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia a.
scales present along anterior margin of scutum from end of submedian line
to dorsal posterior pronotal scales; scutellum with sparse narrow dark
scales on lateral lobes, occasionally pale scales occurring there, mid lobe
with broadened appressed dark scales laterally, a median longitudinal area
of narrower pale scales (extent of this area variable); pleuron with a patch
of broad white scales on each of following areas: postspiracular, lower pre-
alar (hereafter called the "prealar scale patch"), upper sternopleural, lower
posterior sternopleural, and upper half or more of mesepimeral, some spe-
cies also have scales on paratergal and subspiracular areas. Legs. Coxae
with broad white scales, fore and mid coxae also with dark scales ventrally
and fore usually with some white scales antero-ventrad to dark scales; fore
and mid femora with anterior surface dark except often for a narrow fringe
of pale scales basally, pale scales variably present elsewhere; posterior
surface of fore femur with a broad dorsal white band from near base to apex,
may be interrupted or diminished towards apex; posterior surface of mid
femur with basal half largely white, this white continued along apical half as
a ventral band; anterior surface of hind femur with a broad white area from
near base to near middle, connecting ventrally with a similar area on poste-
rior surface (actual location and extent of this area variable with species),
dorsal surface usually dark along whole length, apex white beneath, this ex-
tending apico-dorsally a variable amount onto both anterior and posterior
surfaces; tibiae dark, with a baso-ventral (baso-anterior in some cases)
white area, posterior surface of fore and mid tibiae lined with pale scaling
in some species; tarsi with basal white bands present on some segments;
tarsal claws equal, fore and mid each with 1 tooth, hind simple. Wing.
Dark scaled, nearly always a ventro-basal line of white scales on costa.
Halter. Knob with dark and pale scaling. Abdomen. Terga with dorsal pale
markings usually present; prominent basolateral white spots present, lateral
margin of I with silvery-white scaling; sterna dark scaled, most with basal
white bands.
MALE. In general, similar to female. Head. Palpus about 0. 75-
0.80 length of proboscis, segments straight, not noticeably swollen or
curved, nearly hairless except for a few bristles at apices of segments HI- V
(may also be a very few laterally along the shafts of segments IV-V in some
species, up to about 7 lateral hairs on each side of IV and about 4 on each
side of V); vertex with upright forked scales pale medially, dusky laterally,
sometimes all pale except on submedian dark-scaled areas. Legs. Fore
and mid tarsal claws unequal, the larger claw with a normal median tooth
and a finer acute latero-basal tooth, smaller claw with a sub-basal tooth;
hind claws equal, simple. Wing. Dark scaled. Abdomen. Tergum of seg-
ment I with lateral margin white scaled, remainder of terga with prominent
baso-lateral white spots, some segments with dorso-basal pale scaling or
complete basal bands. Terminalia. Segment IX with tergal lobes small but
prominent, each bearing a number of strong setae; sternum with 1-3 long
setae, usually 1-5 smaller setae, and sometimes 1-3 small scales; basimere
long and rather narrow, densely scaled, mesal surface membranous, re-
mainder distinctly sclerotized, dorsal surface without an apical lobe, usually
without a basal lobe, with 30-60 short setae scattered along mesal margin,
4 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
about 7-20 of these somewhat clustered in area where a basal lobe would be
if there were one, 1-5 of the setae along mesal margin more strongly devel-
oped than remainder, lateral and ventral surfaces with numerous very long
setae; claspette composed basally of a pilose stem bearing 1-6 distinct
setae, and apically of a sickle-shaped filament (elliptically tubular in cross
section) which is slightly longer than the stem; distimere simple, bearing
1-2 setae apically; terminal appendage simple, about 0. 20-0. 25 the length
of distimere itself; phallosome with aedeagus simple, short, when normally
expanded apically it is of rather even width throughout; paraproct with a
single apical tooth, 1-4 cercal setae on each side.
PUPA. (Figure 11.) Cephalothovax. Hair 1-C at least twice as
long as hair 2-C; 10-C mesad of 11-C; 11-C greatly elongate, single.
Abdomen. Hair 1-II usually well developed, with 2 or more branches; 2-I
and 3-I approximated; 2-VI well mesad of 1-VI; 3-I-III greatly elongate,
single; 5-IV-VII noticeably elongate, single; 6-VII approximately similar to
9-VII in development, with 2-5 simple or barbed branches which may be
forked; 9-I-VI minute; 9-VII similar to 6 but usually larger and with more
branches; 9-VIII larger than on VII with 3-14 simple or barbed branches
which are usually forked; 1-P elongate, approximately equal to 5-VII in de-
velopment; paddle margins usually with some fine submarginal spiculation
apically, minutely and sparsely fringed on basal half. Minor differences oc-
cur between the species included here but none has been found to be suffi-
ciently great or stable to permit species determination.
LARVA. (Figure 12.) Head. Antenna moderately long and of even
width with sparse minute spiculation; hair 1-A usually inserted just beyond
middle, with 1-5 branches (usually 2-3); head hair 4-C very small, with 2-7
branches; 6-C with 3-9 and 7-C with 3-9; bases of 4-6-C usually in a rather
even transverse line anterior to antennal bases in unflattened heads; 5, 6-C
with stiff branches which are of rather equal length and arranged in a fan-like
manner; median mouth brush hairs with comb-like tips; mentum with 10-14
lateral teeth, first 2-4 more widely spaced. Abdomen. Hair 1 most promi-
nent dorsal hair on IJI-VII; 6-I-II with 1-4 branches (usually double), with
1-5 on II-VI, branches unequal in length (nearly equal in chrysolineatus);
7-I-II with 1-3 branches (usually single); hair 13 most prominent ventral
hair on II-VI, with 1-6 branches on III-V, single on VI, and with 1-4
branches on VII; 1-VIII with 1-5 branches, 2-VIII and 4-VIII single, 3-VII
with 4-10, and 5-VIII with 2-6 branches; comb consisting of a patch of scales;
siphon hair tuft inserted in area of, or just beyond, middle, with 3-7 plumose
branches; pecten consisting of 8-27 dark teeth, each tooth usually slightly
larger than the more proximal one, teeth basad of hair tuft bearing 1-4 ven-
tral denticles; segment X with pilose integument; saddle incomplete, poste-
rior margin clothed with spicules of varying development; ventral brush con-
sisting of 10-13 tufts, normally all arising from grid; gills 1. 5 to 4 times
longer than saddle, approximately equal in length.
DISTRIBUTION. Throughout the Oriental Region, with representa-
tives in the adjacent portion of the Palaearctic Region.
TAXONOMIC DISCUSSION. The grouping of species created by this
definition has more the characteristics of a ''superspecies" than it does of a
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia a
"group" as used by Edwards (1932). The chrysolineatus subgroup is
presently considered to include the following species: chrysolineatus
(Theobald), formosensis Yamada, harveyi (Barraud), japonicus japonicus
(Theobald), japonicus shintienensis Tsai and Lien, jugraensis (Leicester),
koreicus (Edwards), nigrorhynchus Brug, vizali (Banks), saxicola Edwards,
and sherki Knight. All of these species, except koveicus, occur in the
Southeast Asian Region. 3
The species of this subgroup differ so slightly from one another
that any but perfect specimens are frequently difficult to identify. Species
determinations are most satisfactorily made from a combination of larval
and adult characters. Male terminalia and pupae almost totally lack ade-
quate identification characters.
BIOLOGY. Without exception, the larvae of this group of species
occupy "'container-type" habitats, particularly rock holes, tree holes, and
plant axils. On occasion, the immature stages of some species are found
in artificial containers. Several of the species have been taken in human
biting catches and it is likely that all of the members of the subgroup do, at
least on occasion, feed on man.
KEY TO THE SPECIES - MALES AND FEMALES
ay Scutum with median longitudinal pale line not forked
DORTOTICHD COPA) ge ce i eee a a 2
Scutum with median longitudinal pale line forked posteriorly
atpreecielir space (Figure 1) oo ies etek es le hah oe oe 4
2(1). First 4 hind tarsomeres with basal white bands. ..........
Only first 3 hind tarsomeres basally banded. ........... 3
3(2). Prealar scale patch continuous with upper sternopleural scale
patch (Vigo ee le a Sha Bee jugvaensis (p. 19)
Prealar scale patch distinctly separated from upper sternopleural
Soule parch (Fisare 2) gs a ek a rizali (p. 22)
4(1),. ‘Proboseig ayy Geek ecaled ie ee a 5
Proboscis with pale scaling present. .........-6.6.ee268-. 9
5(4). Mid femur with anterior and antero-dorsal surfaces dark apically;
female torus with dark scales only. ..........60.848%
Mid femur with anterior and dorsal surfaces white just before
apex (Figure 2); torus with white scales’). 6 a kc as 7
6(5).
7(5).
8(7).
9(4).
10(9).
Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
Posterior portion of submedian scutal pale line anteriorly
bent along scutal angle to area of lateral line and of
fateral Scutal mare (2)... 4, +. 3. nigrorhynchus (p. 20)
Posterior portion of submedian scutal pale line anteriorly
bent along scutal angle only briefly, well separated from
Hotere We OW, a a saxicola (p. 25)
Hind leg with first 4 tarsomeres basally white banded, may
be a few basal pale scales on the fifth; subspiracular area
with a line of broad white scales; Korea, China, Japan.....
kRoreicus
Hind leg with first 3 tarsomeres basally banded; subspiracular
Sree WiOut Sele ee ee ese ee ee as he es 8
Female median upright forked scales of head mostly pale;
anterior pronotal lobe with some narrow curved scales
G0rsaily Toiwal, WyORYOs. oe ns ee ee te
japonicus shintienensis (p.17)
Female median upright forked scales of head mostly dark;
anterior pronotal lobe with oblique band of broad white
scales: Japan, Cling... 2. oe japonicus japonicus (p.15)
Paratergite and usually subspiracular area each with a small
petcn Of broad white scales (Fioure 2)... ee Sie ce es
formosensis (p. 10)
Paratergite and subspiracular areas without scales....... 10
Female proboscis with ventral pale-scaled area broadly produced
laterally and, to a somewhat lesser extent, dorsally in general
area of middle; male proboscis with narrow white band
ene ee chrysolineatus (p. 8)
Female proboscis with ventral pale scaling not visible, or only
very slightly so, from above; male proboscis with pale
scaling visible at joint laterally or even from above, but not
forming a sharp complete band........ harveyi (p. 13)
(s) nigrorhynchus (p, 21)
KEY TO THE FOURTH STAGE LARVAE
Abdominal hair I-X basally 2-3 Dranched. . 1"... 6 ke kes hue « 2
Abdominal hair 1-X single, occasionally divided sub-basally... 3
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia
2(1). No pecten teeth inserted beyond siphon hair tuft; abdominal
integument not showing distinctive pilosity...........
nigrorhynchus (p. 21)
Three to 6 pecten teeth inserted beyond siphon hair tuft;
abdominal integument heavily pilose. ..... rizali (p. 23)
sherki (p. 28)
3(1). Metathoracic hair 7-T with stubby branches, distinctly
different in shape and development from hair 6-M
(PROUT Bei cde chs we a
Metathoracic hair 7-T with greatly elongate tapered branches,
generally similar in shape and development to hair 6-M
(PU GUT OTe bok ads Dik uae a ae
4(3). Comb consisting of about 15-20 scales; siphon hair tuft with
tip exceeding siphon apex; pecten teeth with 1 well developed
ventral denticle before middle, no teeth beyond base of
siphon hair. tuit (Vieure 8)... 6 « nw li jugraensis (p.19)
Comb consisting of about 50-70 scales; siphon hair tuft with
tips not attaining siphon apex; pecten teeth with 1-2 very
small ventral denticles medially or with none, 3-6 teeth
inserted beyond base of siphon hair tuft (Figure 10) ......
saxicola (p. 26)
5(3). None of the branches of the siphon hair tuft attaining apex
OL BION De MAINES on, co piss acces pie) Sais bale es
Some, at least, of these branches reaching to or beyond apex
O5 SU NON Ae a OG ie i lp ere ee i oe ah et
6(5). Head hair 5-C with 9-22 branches; comb scales with fine
Spicules basally, coarser ones apically, usually 1 apical
Spicule more strongly developed; siphon blackish, index
about 1.5-2.0, with 8-13 pecten teeth (Figure 4)........
formosensis (p.11)
Head hair 5-C with 3-7 branches; comb scales with fringing
spicules rather even in development; siphon not blackish,
index approximately 3.0-3.5, with 15-29 pecten teeth. ....
7(6). No simple pecten teeth beyond base of siphon hair tuft;
saddle with mostly spiculate scales on posterior margin...
koreicus
Usually 1 or more enlarged simple teeth beyond base of
siphon hair tuft; saddle with mostly simple spines on
posterior margin (Figure 6). . . . japonicus japonicus (p. 16)
japonicus shintienensis (p. 18)
8 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
8(5). Individual comb scales expanded apically; denticles of
pecten teeth small (Figure 5)......... harveyi (p. 13)
Individual comb scales distinctly tapered to a stout
central spine; denticles of pecten teeth large, elongate,
conspicuous (Figure 3)......... chrysolineatus (p. 9)
AEDES (FINLAYA) CHRYSOLINEATUS (THEOBALD)
(Figures 1, 2, 3)
Hulecoeteomyia trilineata Leicester 1904 (non Theobald 1901), im Theobald,
Entomologist 37: 163 (o,?); Theobald 1907, Mon. Cul. 4: 220 (c, 9);
Edwards 1922, Indian J. med. Res. 10: 466 (synonymy).
Howardina chrysolineata Theobald 1907, Mon. Cul. 4: 218 (9).
Culex ? japonicus var. ceylonica Theobald 1910, Mon. Cul. 5: 391 (9);
Edwards 1932, in Wytsman, Genera Insect. fasc. 194: 151 (synonymy)
Aedes (Finlaya) chrysolineatus (Theobald), Borel 1930, Les Moustiques,
248 (o*,9, L*); Barraud 1934, Fauna Brit. India 5: 185 (o*, 9*, L*);
Knight 1947 (1948), Ann. ent. Soc. Amer. 40: 636 (c'*, 2, L).
FEMALE. Head. Torus with dark scaling mesally; palpus broadly
white tipped; proboscis ventrally with extensive pale scaling, this pale
scaling produced onto lateral and dorsal aspect in area of middle as a wide
band, dorsal midline may be dark or band may be complete, exact amount
of pale scaling on proboscis subject to considerable variation; vertex with
upright forked scales dark. Thovax. (Figure 1.) Anterior pronotum with
white scales which are mostly broad; posterior pronotum with many broad
white scales posteriorly, there may also be some present anteriorly and
sometimes there are pale narrow scales dorsally; scutum with pale scaling
yellow to whitish-yellow, median scutal line forked at prescutellar space,
submedian line broken at level of scutal angle with its posterior portion bent
briefly antero-laterally along scutal angle, usually not closely approaching —
lateral scutal margin; pleuron with prealar scale patch continuous with upper
sternopleural patch and no scales on paratergite or subspiracular area.
Legs. (Figure 2.) Fore femur anteriorly dark, a pale-scaled edging ven-
trally; mid femur with pale scales along anterior midline; hind femur with
anterior submedial pale area not crossing dorsal surface, apico-ventral
pale area extensively produced onto anterior surface but not ringing femoral
apex; fore and mid tarsi with distinct basal white bands on I-II, less distinct
basal pale scaling usually present on III of mid tarsus; hind tarsus with
prominent white basal bands on I-III, that on HI dorsally occupying approxi-
mately half of segment. Abdomen. Terga dorsally dark scaled, with nar-
row medio-basal yellowish area or basal bands of white scales usually pres-
ent on some segments (absent on type of chrysolineatus), baso-lateral sil-
very white spots usually visible dorsally on more terminal segments.
Variation. According to Barraud (1934: 188), Malayan specimens differ
from those from India and Ceylon in having few or no white scales on ante-
rior surfaces of fore and mid femora, and apparently no basal bands on
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 9
abdominal terga in female. In scaling of proboscis, posterior pronotum,
etc. , they are closely similar to Indian specimens.
MALE. Head, Palpus with pale scaling at joints of I-IV and IV-V,
remaining portions dark scaled; proboscis dark scaled except for a distinct
white band just beyond middle, additional pale scaling variably present ven-
trally. Thorax. Posterior pronotum with fewer broad scales than in fe-
male. Legs. No pale basal scales on III of fore tarsus. Abdomen. Terga
with dorso-basal pale scaling highly variable, usually more developed than
infemale. Terminalia. (Figure 3.) Tergal lobes of IX each with approxi-
mately 14-34 setae; basimere with a slightly-defined basal tergo-mesal
lobe, bearing 7-28 slender setae.
PUPA. Chaetotaxy similar to saxicola (Figure 11).
LARVA. (Figure 3.) Head. Hairs 5,6-C with 4-6 branches; 14-C
with 2-3 branches; basal maxillary hair (bmh) with 3-4 branches. Thorax.
Without obvious pilosity; hair 1-P with 2-3 branches; 3-P with 4-6; 8-P with
4-6; 14-P with 2-3; 1-M with 3-4; 1-T with 3-5; 5-T with 1-2; 7-T with 4-6,
Similar in shape and development to hair 6-M. Abdomen. Pilosity, if pres-
ent, small and not uniformly distributed; hair 1-II-VII with 2-4 branches;
comb consisting of 22-27 scales, each scale short, sharply tapered toa
stout central spine, with lateral fringe of short spicules (none of which at-
tains apex of central spine); siphon non-pilose, pale, index approximately
2. 2-2. 5; siphon hair tuft with tips usually reaching or slightly exceeding
siphon apex; pecten with 13-19 teeth, with 2-4 distinct ventral denticles,
last 1-2 teeth usually inserted on a level with or just beyond siphon hair tuft,
terminal tooth may be simple or with denticles; hair 1-X stout, barbed,
single, considerably longer than saddle; ventral brush without precratal
tufts. Variation. This description, which has been prepared from Thailand
specimens only, matches all details given for the larva of chrysolineatus by
Barraud (1934: 187; presumably based upon Indian specimens) except that he
describes hair 1-X as being 3-4 branched and only a little longer than the
saddle. Viet Nam material as described by Borel (1928, 1930) resembles
that from Thailand in having this hair single.
TYPE DATA. Hulecoeteomyia trilineata Leicester, cotype male and
female in British Museum, male terminalia mounted; type locality: Kuala
Lumpur (Leicester), MALAYSIA. Howardina chrysolineata Theobald, holo-
type female in British Museum; type locality: Pundabroya (= Pundaluoya;
Green), CEYLON. Culex japonicus var. ceylonica Theobald, 1 cotype fe-
male in British Museum, location of second cotype unknown; type locality:
Peradeniya (Green), CEYLON.
DISTRIBUTION. Specimens examined: INDIA, Bombay, Nagargali
and Deccan, 3 males, 3 females. THAILAND, Chiang Mai, Chumphon,
Ranong, and Nan, 25 males, 11 females, 2 pupae, 39 larvae, 10 rearings.
VIET NAM, Quan Loi, 6 larvae. Records from literature: INDIA, Bombay,
Malabar, Eastern Himalayas. CEYLON (Barraud, 1934). THAILAND,
Patalung Pass (Causey 1937). MALAYSIA, Bukit Kutu and Ulu Gombak, nr.
Kuala Lumpur (Leicester 1908). JAVA, Ngledok (Brug 1931), Tosari
(Theobald 1911). VIET NAM (as Cochinchina, Borel 1928). SUMATRA
(Brug and Bonne-Wepster 194’).
10 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
TAXONOMIC DISCUSSION. Because of the variation in type and
extent of adult markings which occurs in this species, it is frequently dif-
ficult to separate from harveyi, which is not only very similar morphologi-
cally but also on occasion occurs in the same larval habitat. Fortunately,
the larvae are sharply distinct on the structure of the comb scales and
pecten teeth. Accordingly, on the basis of a recently-acquired series of
reared adult-associated larval and pupal skins from Thailand, it is now pos-
sible to describe the range of adult variation for the 2 species with some
measure of confidence. It is likely that the Thailand material will eventu-
ally prove to be a species other than chrysolineatus for Barraud (1934:185)
describes Indian larvae as having hair 1-X with 3-4 branches, whereas it is
uniformly single in larvae from Viet Nam (Borel 1928, 1930) and from
Thailand. This is a character of specific rank in the chrysolineatus sub-
group. Adult-associated rearings from India will be required for the solu-
tion of this problem. ,
BIOLOGY. The larva has been reported from tree holes (Barraud
1923), from rock holes in a mountain stream (Causey 1937), from bamboo
(Leicester 1908; Barraud 1924), from Colocasia leaf axils (Brug 1931), and
from roof gutters and broken chatti (Barraud 1934). The Thailand speci-
mens (USNM) newly reported here were reared from the following habitats:
fallen leaf, banana tree leaf axil, rock pool, root hole, bamboo stump, bam-
boo internode, tree hole, anda clay pot. The Viet Nam specimens were
from rubber cups. Leicester (1908) and Macdonald (1957) recorded the
adult as being a sylvan biter of man.
AEDES (FINLAYA) FORMOSENSIS. YAMADA
(Figures 1, 2, 4)
Aedes formosensis Yamada 1921, Annot. zool. jap. 10:67 (¢).
Aedes (Finlaya) pallirostris Edwards 1922, Indian J. med. Res. 10: 270 (¢);
Barraud 1934, Fauna Brit. India 5: 190 (?); Knight 1947 (1948), Ann.
ent. Soc. Amer. 40: 640 (?). NEW SYNONYMY. a
Aedes formosaensis Yamada, Edwards 1922, Indian J. med. Res. 10: 262
(lapsus).
Finlaya khasiana Barraud 1923, Bull. ent. Res. 13: 407 (c, ?); Edwards
1932, in Wytsman, Genera Insect. fasc. 194: 151 (synonymy).
FEMALE. Head. Torus with dark scales mesally; palpus broadly
white tipped, sometimes a few pale scales at apex of penultimate segment;
proboscis pale scaled beneath in middle third, the pale scaling visible from
lateral aspect, not usually visible from above ("Rkhasiana" form); exact ex-
tent of pale scaling subject to much variation, in some specimens pale scaling
encroaches onto dorsal surface and may on occasion even cover all but mid-
line of basal four-fifths of proboscis ("pallirostris" form); vertex with medi-
al upright forked scales dark. Thorax. Anterior pronotum with broad
white scales; posterior pronotum with narrow curved yellowish or whitish
scales, at least some broad pale scales ventro-posteriorly; scutum
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 11
(Figure 1) with median line forked at prescutellar space; submedian line
strongly developed, broken at level of scutal angle with its posterior portion
bent antero-laterally along scutal angle to lateral margin; line over wing
base anteriorly closing or nearly so at scutal angle with outcurved portion of
submedian line; pleuron (Figure 2) with scales noticeably broad in compari-
son with most of the species of this subgroup; prealar scale patch distinctly
separate from upper sternopleural scale patch; paratergite with a line of
broad white scales, mostly borne on lower surface; subspiracular area often
with a small group of broad white scales. Legs. Fore femur anteriorly
dark, a pale-scaled line along ventral border and a small poorly defined
creamy area dorsally at apex; mid femur usually with scattered (may be
continuous) pale scaling along part or all of longitudinal midline anteriorly,
usually a small white spot dorsally just before apex; hind femur with apico-
ventral pale area extensively produced onto anterior surface and usually en-
circling apex; tibiae each with very pronounced basal white area anteriorly
and a thin white basal ring; fore and mid tarsi with distinct basal white bands
on tarsomeres I and II, less distinct basal pale scaling usually present on
mid tarsomere III; hind tarsus with prominent white basal bands on tarso-
meres I-III, that on III dorsally occupying approximately half of segment.
Abdomen. Terga dorsally dark scaled, narrow medio-basal areas of pale
scales (usually of an obscure yellowish-white color but may be bright white)
present on some segments (usually III-VI), basolateral silvery-white spots
usually visible dorsally on more terminal segments. Variation. A female
from Sumatra has the proboscis so dark scaled that it was first believed to
so into that portion of the adult key. However, careful examination showed
the proboscis to have a small ventral area, just beyond the middle, of pale
brownish scales.
MALE. Head. Palpus with basal pale scaling ventrally and laterally
on IV and V, a prominent basal white band laterally and dorsally on III; pro-
boscis with a narrow area of pale scales ventrally and laterally at joint, may
be visible dorsally, additional pale scaling may variably occur along ventral
surface. Abdomen. Terga with basal white bands on II-VII, these may be
disconnected laterally from baso-lateral silvery-white spots on 1 or more
segments, II may have medio-basal pale scales. Terminalia. (Figure 4.)
Tergal lobes of IX each with approximately 4-13 setae; claspette filament
not noticeably wider at any particular point, tapering to apex from about
middle. Variation. A male (USNM: CM-148) from Thailand has the pro-
boscis completely dark scaled. This character is of specific rank in this
subgroup, but it never applies quite as well to the male as to the female.
Its significance in this particular case is not known.
PUPA. Chaetotaxy similar to saxicola (Figure 11).
LARVA. (Figure 4.) Head. Hair 5-C with 9-22 branches; 6-C with
5-9; 14-C and basal maxillary hair (bmh) stiff, single, rarely double.
Thorax. Without obvious pilosity; hair 1-P with 2-7 branches; 3-P with 8-
15; 8-P with 3-6; 14-P with 2-3; 1-M well developed, with 4-8 branches; 1-T
with 2-6 branches; 5-T stiff, single; 7-T with 4-12, similar in form and less
prominently developed than hair 6-M. Abdomen. Pilosity, if present, very
small and not uniformly distributed; hair 1-III-VII with 2-7 branches; comb
12 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
consisting of 29-51 scales, each with fine lateral spicules basally and
coarser spicules along remainder of length, usually 1 apical spicule much
more strongly developed; siphon non-pilose, blackish-brown, index approx-
imately 1. 5-2. 0; siphon hair tuft with tips not attaining siphon apex; pecten
with 8-13 teeth, each tooth with 1-2 ventral denticles along basal half, may
be 1-3 non-dentate teeth with point of attachment even with or beyond in-
sertion of siphon hair tuft; hair 1-X stout, barbed, single, length about 1.5
times length of saddle; ventral brush with 1-2 precratal tufts.
TYPE DATA. Aedes formosensis Yamada, female lectotype (se- :
lected by M. Delfinado, VIII-1966) in Medical Zoology Laboratory, Institute
for Infectious Diseases, University of Tokyo; type locality: Kakubanzan
(given as 'Kappanzan, Formosa" on label; Hirayama), TAIWAN. Aedes
pallirostris Edwards, female holotype in British Museum; Golaghat,
Sibsagar dist., Assam (Christophers), INDIA. Finlaya khasiana Barraud,
male and female cotypes in British Museum; type locality: Shillong, Khasi
Hills, Assam (Barraud), INDIA.
DISTRIBUTION. Specimens examined: MALAYSIA, 4 females.
THAILAND, Chiang Mai, Narathiwat, Khaoyai, Pakchong, Nan, Nakhon Si
Thammarat, 32 males, 77 females, 52 larvae, 26 rearings. SUMATRA,
Ranau-See, 1 male, 2 females. JAVA, Tijsaroea, Patjat, Ngledok,
Tjisanaea,1 male, 6 females. BALI, Batoeriti, 1 female. Records from literature:
TAIWAN (Yamada 1921, Lien 1962). INDIA, Assam, E. Himalayas (Barraud
1934). ?SUMATRA. ?BALI (Edwards, in Barraud 1934). CHINA, Western
Yunnan (Chow 1949). MALAYSIA, Ulu Langat, Ulu Gombak, Kepong
(Selangor) (Macdonald 1957).
TAXONOMIC DISCUSSION. Based upon an examination of the type of
formosensis by M: Delfinado (August, 1966), pallirostris is here synony-
mized to that species. Edwards (in Barraud 1934) earlier suggested that
pallirostris might in fact be just a variation of formosensis, but an examina-
tion of the type of the latter was required to confirm this.
Most of the specimens included here represent the form previously
treated by Knight (1947) as formosensis because they are similar to the type
of khasiana in having little or no pale scaling dorsally on the proboscis.
Specimens with distinct dorsal pale scaling on the proboscis were considered
to be pallirostris because its type is similarly marked.
A long series of specimens from Thailand have recently become
available, some with associated larval and pupal skins. From an examina-
tion of these specimens, it is now apparent that the extent of the pale scaling
of the proboscis is subject to much variation, and that both extremes can be
found associated with a single larval type.
A. formosensis is unique in the chvysolineatus subgroup in possessing
scales on the paratergite of the adult, and in having larval head hair 5-C vir-
tually plumose (with 9-22 branches).
BIOLOGY. In Thailand, this species was a frequent component of
daytime biting catches, primarily in the forest, but occasionally also in the
open. The larvae were collected commonly in the leaf axils of Colocasia and
of banana trees. One collection was from a bamboo stump. Lien (1962)
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 13
reported that in Taiwan the larvae of this species were found almost
exclusively in leaf axils of Musa and more rarely in Colocasia.
AEDES (FINLAYA) HARVEYI (BARRAUD)
(Figure 5)
Finlaya harveyi Barraud 1923, Bull. ent. Res. 13: 407 (¢, 2).
Aedes (Finlaya) harveyi (Barraud), Barraud 1934, Fauna Brit. India 5: 188
(o, 2); Knight 1947 (1948), Ann. ent. Soc. Amer. 40: 638 (, 9).
Aedes (Finlaya) formosensis Yamada, Chow 1950, Quart. J. Taiwan Mus. 3:
282 (misidentification); Lien 1962, Pacif. Ins. 4: 622 (in part).
FEMALE. Head. Torus with dark scaling mesally; palpus white
tipped; proboscis with pale scaling beneath from near base to shortly beyond
middle, dark laterally and dorsally, actual extent of ventral pale scaling
variable, may even be some extension onto lateral surface; vertex with up-
right forked scales dark. Thorax, Anterior pronotum with white scales,
mostly broad; posterior pronotum with narrow whitish or yellowish scales
above, a small patch of broad white scales ventro-posteriorly; scutal pale
scaling golden, median scutal line forked posteriorly at prescutellar space,
submedian line broken at level of scutal angle with its posterior portion bent
antero-laterally along scutal angle, laterally connecting or nearly so with
anterior portion of lateral line; prealar scale patch usually continuous with
upper sternopleural scale patch, sometimes showing evidence of separation,
occasionally completely separated, no scales on paratergite or subspiracular
area. Legs. Fore femur anteriorly dark, marked with a pale scaled line
ventrally along basal half; mid femur with anterior surface usually dark;
hind femur with anterior submedial pale area not extending onto dorsal sur-
face, apico-ventral white area broadly produced onto anterior surface but
not quite ringing apex; fore and mid tarsi with distinct basal white areas on
I-II, less distinct basal pale scaling usually present on middle tarsal III;
hind tarsus with prominent white basal bands on I-III, that on III dorsally oc-
cupying slightly more than half of segment. Abdomen. Terga without dorsal
markings, baso-lateral silvery spots usually visible dorsally on more ter-
minal segments, occasionally a few baso-medial yellowish scales occur on
some terga.
MALE. Head. Palpus with basal pale scaling ventrally and laterally
at joints of III-IV and IV-V; proboscis with a narrow area of pale scales ven-
trally and laterally at joint, may extend very narrowly onto dorsal surface.
Abdomen. Atleast some terga, usually III-VI, with pale basal bands. Ter-
minalia, (Figure 5.) Tergal lobes of IX each with approximately 13-26
setae; basimere with a slightly-defined basal tergo-mesal lobe, bearing 7-15
slender setae; claspette filament may be slightly enlarged in area of middle,
the taper beginning at or just beyond middle.
PUPA. Chaetotaxy similar to saxicola (Figure 11).
LARVA. (Figure 5.) Head. Hairs 5,6-C with 4-7 branches; 14-C
and basal maxillary hair (bmh) well developed, former with 1-4 (usually 1-2),
14 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
latter with 3-5. Thovax. Without obvious pilosity; hair 1-P with 2-4
branches; 3-P with 2-6; 8-P with 4-9; 14-P with 2-3; 1-M well developed,
with 3-6 stiff stubby branches; 1-T with 3-6 branches; 5-T with 1-2; 7-T
with 3-9, similar in shape and development to 6-M. Abdomen. Pilosity if
present very small and not uniformly distributed; hair 1-III-VII with 1-3
branches; comb consisting of 26-46 scales, each scale slightly expanded api-
cally and with an even lateral and apical fringe of spicules; siphon non-pilose,
index approximately 2. 4-2. 7; siphon hair tuft with tips usually reaching or
slightly exceeding siphon apex; pecten with 12-21 teeth, with 1-4 ventral
denticles on basal half of all except terminal 1-2 teeth (rarely none without
denticles), last tooth usually inserted just before base of siphon hair tuft,
occasionally 1 non-dentate tooth with point of attachment beyond insertion of
siphon hair tuft; hair 1-X stout, barbed, considerably longer than saddle,
Single (occasionally seen sub- -basally divided): ventral brush without precra-
tal tufts.
TYPE DATA. Male and female cotypes in British Museum; type
locality: Kurseong, Darjeeling district, Eastern Himalayas (Barraud),
INDIA.
DISTRIBUTION. Specimens examined: VIET NAM, Dalat, 2 males,
3 females, 3 rearings. MALAYSIA, Selangor, 3 males, 5 females, 7 rear-
ings. INDIA, Kurseong, Eastern Himalayas, 1 male, 1 female. THAILAND,
Chiang Mai, Nakhon Nayok, Nakhon Si Thammarat, Chanthaburi, 23 males,
42 females, 153 larvae, 51 rearings. TAIWAN, 4 females. SUMATRA,
Tjoeroep, 1 male. Records from literature: INDIA, Eastern Himalayas,
Madras, South India (Barraud 1934). SUMATRA, Lake Banau, 700 m.
Boekit Daoen (Brug and Edwards 1931). ?JAVA, Negadisari and Tjibodas.
?BALI, Batoeriti and Kintamani (Brug 1931). CEYLON (Carter and
Wijesundara 1948). MALAYSIA (Macdonald 1957). CHINA, Western Yunnan
(Chow 1949). |
TAXONOMIC DISCUSSION. The adult of harveyi is so similar to
chrysolineatus that the 2 species are frequently difficult to separate in that
stage. Although the types had been seen earlier, the concept of harveyi used
here was not established until adult-associated larval skins from Thailand
and Malaysia were studied. Based upon adult-associated larval skins of both
Species, it is now possible to say that they are quite distinct on the basis of
larval comb and pecten tooth structure. Just as with chrysolineatus, the pos-
sibility exists that, when adult-associated larvae are seen from India, they
will prove to be different from those from Thailand and Malaysia, thereby
making the latter an unnamed species.
The larva of harveyi was previously depen bed by Brug (1931) from
a series of larvae and adult-associated larval skins collected in Sumatra,
Java, and Bali. As described by him, the Indonesian larva differs markedly
from the Malaysian and Thailand material described above, as follows: head
hair 5-C figured with 9 branches; comb scales of 2 types with posterior
scales illustrated as being sharply tapered apically, and pecten shown to
have 2 strong teeth well distad of the insertion of the siphon hair tuft. I have
not seen any of Brug's specimens so cannot accurately determine what
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 15
species he had. This larva will go to the harveyi-chrysolineatus couplet in
the larval key of this paper, but is distinct from either of these in having
apical pecten teeth and 2 types of comb scales.
Edwards (in Barraud 1934:190) subsequently saw 2 of Brug's isola-
tions (from Bali) and pronounced them formosensis. His description of
these 2 larvae does indeed sound like formosensis. However, the larva
figured by Brug (1931) does not match my concept of formosensis since the
siphon hair tuft reaches the siphon apex.
The identity of these various specimens will remain in doubt until
adequate series of adult-associated larvae are available from Indonesia and
India.
Four female specimens from Taiwan have been seen in the USNM
- collection which are probably harveyi. In each case they were reared from
bamboo stumps. Lien (1962) says that although Chow (1950) recorded
formosensis from bamboo stumps, he had not been able to collect it from
this habitat; rather all specimens resulting from larvae collected from bam-
boo stumps were another species in the chrysolineatus group, which he plan-
ned to discuss in a separate paper. I have not yet seen that paper but I think
it likely that his species from bamboo and harveyi are one and the same.
BIOLOGY. Because of the uncertainties existing in the identification
of this species, the larval habitats recorded by Brug (1931) are not given
here. In Thailand, larvae which I have examined were collected from the
following habitats: cavity on top of a log, in a clay pot, tree hole, bamboo
stump, fallen leaf, rock pool, coconut shell, bamboo internode, and tin con-
tainer. On at least 1 occasion, an adult was taken in a biting collection in
heavy forest and a male was found resting in a hut. The type series was
collected from tree holes.
AEDES (FINLAYA) JAPONICUS JAPONICUS (THEOBALD)
(Figure 6)
Culex japonicus Theobald 1901, Mon. Cul. 1: 385 (9).
Aedes (Finlaya) eucleptes Dyar 1921, Insec. Inscit. menst. 9: 147 (¢, ?);
Edwards 1922, Indian J. med. Res. 10: 465 (synonymy).
Aedes (Finlaya) japonicus (Theobald), Knight 1947 (1948), Ann. ent. Soc.
Amer. 40: 631 (¢, 9, L*); LaCasse and Yamaguti 1950, Mosq. Fauna
Japan and Korea, 151 (o*, 9*, P*, L*),
FEMALE. Head. Torus with mesal aspect bearing both dark and
pale broadened scales; palpus dark scaled; proboscis dark scaled; vertex
with upright forked scales dark, sometimes a few pale ones medially.
Thorax. Anterior pronotum with an oblique band of broad white scales; pos-
terior pronotal scaling white, yellowish, or mixed, considerable variation
exists in the arrangement, amount, extent, and precise shape of the scales;
scutum with golden scales longer than brown scales, having a roughened or
untidy appearance; median scutal line broad, sometimes tending to appear
double, forked posteriorly at prescutellar bare space; submedian line broken
16 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
at level of scutal angle with anterior end of posterior portion bent laterally
along scutal angle, sometimes extending as far as lateral margin of scutum
at anterior end of lateral line, posterior end of anterior portion of submedian
line bent slightly mesad so as to pass briefly inside of posterior portion; a
broad diffused pale scaled area laterally over wing base; in some specimens
part or all of scutal lines become obliterated by diffused golden scaling;
pleuron with prealar and upper sternopleural scale patches separated and no
scales on paratergite or subspiracular area. Legs. Fore femur dark ante-
riorly, sometimes a baso-ventral line of white scaling; mid femur dark an-
teriorly except for a dorsal white area almost at apex (there is an apical fringe
of dark scales),some pale scales may occur along longitudinal midline;hind
femur with the median anterior white area occasionally continuous, at least
narrowly, to base, the basal extent of this marking apparently subject to
much variation; apical pale area forming a complete ring almost at apex (an
apical fringe of dark scales is present); fore tarsus with basal white scaling
on I-II; mid tarsus with distinct basal white bands on I-II; hind tarsus with
prominent basal white bands on I-HI, that on NI occupying slightly less than
half of the segment. Abdomen. Terga dorsally dark scaled; baso-lateral
silvery-white spots usually visible dorsally on VII-VIII, sometimes also on
IV-VI; may be a few medio-basal pale scales on some segments.
MALE. Head. Palpus dark scaled; proboscis dark scaled. Thorax.
Scutum with golden markings even more diffused and roughened than in female.
Abdomen. Terga dorsally dark scaled, with a baso-medial spot of pale
scales on II-VI, may be reduced to just a few scales on some segments,
baso-medial spots may occasionally form narrow basal bands by connecting
laterally with baso-lateral spots. Terminalia. (Figure 6.) Tergal lobes of
IX each with approximately 4-7 setae; tergo-mesal margin of basimere me-
dially with 1 seta noticeably more strongly developed than the others, and in
this respect differing from other members of the subgroup; claspette fila-
ment evenly tapered.
PUPA. No specimens available.
LARVA. (Figure 6.) Head. Hairs 5,6-C with 3-6 branches; 14-C
and basal maxillary hair (bmh) stiff, single or rarely double. Thorax. With-
out obvious pilosity; hair 1-P with 2-3 branches; 3-P with 2-5; 8-P with 1-4;
14-P with 2; 1-M with 1-5; 1-T small, with 2-4 branches; 5-T single; 7-T
with 4-7, similar in shape and development to 6-M. Abdomen. Pilosity, if
present, very small and not uniformly distributed; hair 1-II-VII with 1-3
branches; comb consisting of about 38-74 scales; each tooth slightly enlarged
apically and bearing a rather even lateral and apical fringe; siphon non-pilose,
index approximately 3. 0-3. 5; siphon hair tuft with tips not attaining siphon
apex; pecten with 15-27 teeth, with 1-5 ventral denticles along basal half of
each tooth, usually 1-4 irregularly spaced non-denticulate teeth beyond base
of siphon hair tuft; hair 1-X 1.0-1.5 times length of saddle, not heavily
barbed, single; ventral brush with 1-2 precratal tufts.
TYPE DATA. Culex japonicus Theobald, 2 female cotypes in British
Museum; type locality: Tokyo (Woods), JAPAN. Aedes eucleptes Dyar, male
holotype in U. S. National Museum, terminalia mounted; type locality:
Canton (Howard), CHINA.
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 17
DISTRIBUTION. Specimens examined: JAPAN, Honshu, Kyushu,
17 males, 16 females, 35 larvae. CHINA, Foochow, 2 males, 2 females,
10 larvae. HONG KONG, 1 larva. Records from literature: JAPAN,
Honshu, Shikoku, Kyushu (Yamada 1927); Hokkaido (LaCasse and Yamaguti
1950). CHINA, Anhwei, Chekiang, Fukien, Kiangsi, Kwangtung (Feng 1938).
SOVIET FAR EAST (Chagin and Kondratiev 1943).
TAXONOMIC DISCUSSION. This species has been commonly confused
with Roreicus. However, it is well distinct from it on adult and larval char-
acters. The Taiwan material formerly attributed to this species is now be-
ing considered under the subspecies, japonicus shintienensis Tsai and Lien,
as is also the southern Ryukyus material of Bohart (1959).
BIOLOGY. The larva has been reported from a variety of container
habitats, including granite cemetary basins, stream rock holes, cement
tanks, bamboo stumps, tree holes, and artificial containers near homes.
The larva is occasionally collected from ground pools. The female bites
humans but only uncommonly.
According to Yamada (1927), japonicus is not a suitable intermediate
host of W. bancrofti. It has been reported as a vector of Japanese encepha-
litis in the Soviet Far East by Chagin and Kondratiev (1943).
AEDES (FINLAYA) JAPONICUS SHINTIENENSIS TSAI AND LIEN
(Figures 1, 2, 7)
Aedes (Finlaya) shintienensis Tsai and Lien 1950, J. med. Ass. Formosa
49: 177 (o*, 9*, P*, L*),
Aedes (Finlaya) japonicus shintienensis Tsai and Lien, Lien 1962, Pacif.
Ins. 4: 623.
FEMALE. Head. Torus with mesal aspect bearing both dark and
pale broadened scales; palpus dark scaled; proboscis dark scaled; vertex
with broad medial area of pale upright forked scales. Thorax. Anterior
pronotum with narrowed yellowish-white scales above, broad white scales
below; posterior pronotum with narrow-curved yellowish-white scales ante-
riorly, dorsally, and posteriorly, a small patch of broad whitish scales
ventro-posteriorly; scutum (Figure 1) with golden scales appearing longer
and more shaggily arranged than the brown scales; median scutal line broad,
somewhat double in appearance, forked posteriorly at prescutellar bare
Space; submedian line broken at level of scutal angle, anterior portion mesal-
ly out of line with posterior portion, anterior end of posterior portion bent
laterally along scutal angle and vaguely connecting with anterior end of later-
al line; area between submedian and lateral lines with diffused golden scaling;
prealar and upper sternopleural scale patches separate; no scales on para-
tergite or subspiracular area. Legs, Fore femur dark anteriorly, some-
times a baso-ventral line of pale scaling; mid femur dark anteriorly except
for a small basal white area, a dorso-subapical white area (a single ring of
apical black scales), and usually some pale scales along longitudinal mid-
line (Figure 2); hind femur with broad anterior white area extending from
18 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
near base to beyond middle, apical pale area forming a complete ring (in
Figure 2 , however, an apical fringe of dark scales is present); tibiae dor-
sally and anteriorly dark; fore and mid tarsi with narrow basal white bands
on I-II; hind tarsus with prominent basal white bands on I-III. Abdomen.
Terga dorsally dark, baso-lateral silvery-white spots visible dorsally on
more terminal segments.
MALE. Head. Palpus dark scaled; proboscis dark scaled. Thorax.
In some specimens, scutum has diffused golden scales overall with an obli-
teration of linear pattern. Abdomen. Terga dorsally dark scaled, witha
baso-medial spot of pale scales on III-VI, these spots may be reduced to just
a few scales on some segments, or even absent. Terminalia. Tergal lobes
of IX each with approximately 5-6 setae; tergo-mesal margin of basimere
with 5-6 setae more strongly developed than those adjacent; claspette fila-
ment slightly enlarged just before middle.
PUPA. No specimens available (figured but not described in type
description).
LARVA. (Figure 7.) No adequately preserved larvae from Taiwan
were available. However, from the type description the larva does not ap-
pear to differ from that of 7. japonicus. The ventral brush is described as
having only 9 tufts, whereas in j. japonicus it has 11-12 tufts. However,
this difference is possibly due to an error in observation. Two larvae from
Iriomote in the Ryukyu Islands (USNM) did not show any noticeable differ -
ences from Japanese larvae of j. japonicus except that in the former the last
1-2 pecten teeth were slightly out of line ventrally. One of these specimens
has been figured.
TYPE DATA. Cotype male and female presumed to be in Inst. Trop.
Med. , National Taiwan University, Taipeh, Taiwan; type locality: Chtichih,
Shintien, Taipeh (Lien), TAIWAN.
DISTRIBUTION. Specimens examined: TAIWAN, 2 males, 12 fe-
males, 6 larvae. RYUKYU ISLANDS, Iriomote Island, Ishigaki Island, 1
male, 1 female, 2 larvae. Records from literature: TAIWAN (Tsai and
Lien 1950).
TAXONOMIC DISCUSSION. This form differs from the type species
in having a large median area of pale upright forked scales on the vertex in
the female, in having narrow-curved yellowish-white scales dorsally on the
anterior pronotum, and in the absence of a single enlarged seta medially on
the tergo-mesal margin of the basimere. Specimens from Japan, apparently
j. japonicus, normally have the upright forked scales dark but specimens
have been seen from Kyushu which had a few of the median upright forked
scales pale.
BIOLOGY. Tsai and Lien (1962) report that the larva of this form is
common throughout Taiwan in rock holes in stream beds, depressions on
fallen trees, and artificial containers in mountainous areas below 2400 me-
ters. The live larva has a dark brown head and reddish brown thorax and
abdomen. The habits of the adult are not known.
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 19
AEDES (FINLAYA) JUGRAENSIS (LEICESTER)
(Figures 1, 2, 8)
Hulecoeteomyia jugraensis Leicester 1908, Cul. Malaya: 109 (c, 9).
Aedes (Finlaya) jugraensis Leicester, Edwards 1922, Indian J. med. Res.
10: 466; Knight 1947 (1948), Ann. ent. Soc. Amer. 40: 647 (c, 2, L);
Knight and Hull 1951, Pacif. Sci. 5: 242 (L*).
FEMALE. Head. Torus with dark scales mesally; palpus apically
white scaled; proboscis dark scaled; vertex with upright forked scales dark.
Thorax. Anterior pronotum with broad creamy scales; posterior pronotum
with a dorsal semicircle of yellowish scaling, narrow curved except fora _
small posterior area of broadened scales; median scutal line (Figure 1) ex-
tends posteriorly unforked nearly or completely across prescutellar area;
submedian line broken at level of scutal angle, sometimes with its posterior
portion bent briefly antero-laterally along the angle; line over wing base an-
teriorly not connecting at scutal angle with outcurved portion of submedian
line; prealar scale patch (Figure 2) confluent with upper sternopleural scale
patch; no scales on paratergite or subspiracular area. Legs. Fore and
mid femur anteriorly dark scaled, a baso-ventral line of white scaling on
fore femur; hind femur with ventro-apical white scaled area extending onto
both anterior and posterior surfaces and narrowly ringing apex (Figure 2);
fore and mid tarsi with narrow basal white rings on I-II, hind tarsus with
broad basal bands on I-III, that on III occupying the basal 0.4-0.5 of the
segment. Abdomen. Terga I-VII dark dorsally, baso-lateral white spot
visible dorsally on IV-VI and extending prominently onto dorsum of VII, VIII
with a complete basal band. Variation. A female specimen from Tjisaroea,
Java (BM), with abdomen missing, matches this species except that it has
the prealar and upper sternopleural scale patches widely separated. In this
respect, it resembles A. vizali, differing from it only in having the posterior
portion of the submedian scutal line not bent antero-laterally along the scutal
angle. The distribution is, of course, far removed from the presently known
range of A. vizali. :
MALE. Head. Palpus dark scaled, terminal segment entirely white
beneath; proboscis dark scaled. Abdomen. Terga IV-VI with basal white
bands, medio-basal white spot on III, baso-lateral silvery spots on VII-VHI
extend onto dorsal surface. Terminalia. Tergal lobes of IX each with ap-
proximately 6-10 setae, basimere with a distinctive band of short curved
setae along basal meso-tergal area; claspette filament also distinctively
shaped,not sickle-shaped but blade-like with widest point at about 0.75 from
base (Figure 8).
PUPA. No specimens available.
LARVA. (Figure 8.) Head. Hairs 5,6-C with 3-4 branches; 14-C
with 1-2; basal maxillary hair (bmh) with 4-8. Thorax. Without obvious
pilosity; hairs 1, 3-P with 3-4 branches; 8-P stellate, with 5-8; 14-P with 2;
1-M dark, spine-like, single; 1-T with 2-3; 5-T stiff, single; 7-T with 3-4
branches, each short, very stout, darkly pigmented, and barbed, very unlike
6-M in shape and development. Abdomen. Without noticeable pilosity; hair
20 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
1-IJI-VII with 2-4 branches; comb with about 14-19 large scales, each scale
with a lateral fringe of evenly-developed spicules and a stout apical spine or
prolongation; siphon non-pilose, index approximately 2.4; siphon hair tuft
with tips exceeding siphon apex; pecten with about 10-16 teeth, each tooth
with 1 well developed ventral denticle before middle and sometimes a second
smaller one basally, no teeth inserted beyond base of siphon hair tuft; hair
1-X single, nearly twice length of saddle; ventral brush without precratal
tufts.
TYPE DATA. Male and female cotypes, non-existent; type locality:
Jugra (Leicester), MALAYSIA.
DISTRIBUTION. Specimens examined: PHILIPPINES, Balabac
Island, 2 larvae. MALAYSIA, Perak and SINGAPORE,1 male, 3 females, 1
larva. N. BORNEO, Tawau, 1 male. JAVA, Tjisaroea, 1 female. Records
from literature: MALAYSIA, ? PHILIPPINES (Macdonald 1957).
TAXONOMIC DISCUSSION. Based upon the larva, this species is
closely related to saxicola, differing principally from that species on the
characters used in the keys. The male is distinct from all other species in
the subgroup on the location of the palpal pale scaling, the shape of the clas-
pette filament, and the setation of the basal tergo-mesal area of the basi-
mere.
BIOLOGY. Macdonald and Traub (1960) reported this to be the most
common member of the chrysolineatus subgroup in Malaysia. Nearly half of
their collections came from fallen split bamboos. Occasional collections of
larvae were also made from bamboo stumps, bamboo sections with holes,
and tree holes, but the most important secondary source of larvae was fallen
leaves. They have also been found in rock pools and in tin cans. The adults
were collected from human bait (Macdonald 1957).
AEDES (FINLAYA) NIGRORHYNCHUS BRUG
Aedes (Finlaya) harveyi var. nigrorhynchus Brug 1931, Arch. Hydrob. Suppl.
bd. 9: 28 (¢%, 9, L*); Knight 1947 (1948), Ann. ent. Soc. Amer. 40: 636
(taxonomy). NEW STATUS.
FEMALE. Head. Torus with dark scales mesally; palpus white tip-
ped, sometimes a few pale scales at apex of penultimate segment also; pro-
boscis all dark scaled; upright forked scales of vertex dark. Thovax. Ante-
rior pronotum with broad white scales; posterior pronotum with narrow and
narrow-curved pale scales dorsally and posteriorly, a few broadened pale
scales ventro-posteriorly; scutal pale scaling golden; median scutal line
forked posteriorly at prescutellar space; submedian line broken at level of
scutal angle with its posterior portion bent antero-laterally along scutal angle,
connecting laterally with anterior portion of lateral line; prealar scale patch
distinctly separate from upper sternopleural scale patch; paratergite and
subspiracular area without scales. Legs. Fore femur anteriorly dark,
marked with a pale-scaled line ventrally along basal half; mid femur with an-
terior surface dark; hind femur dark dorsally, apico-ventral white area
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 21
extensively produced onto anterior surface; fore and mid tarsi with distinct
basal white bands on I-II; hind tarsus with prominent basal white bands on
I-III, that on III dorsally occupying slightly more than half of segment.
Abdomen. Terga dorsally dark scaled, baso-lateral silvery spots usually
visible dorsally on VII-VIII.
MALE. Head. Palpus with basal pale scaling ventrally and laterally
on IV and V; proboscis with a narrow area of pale scales ventrally and later-
ally at joint which may extend onto dorsal surface. Abdomen. At least some
terga with narrow pale basal bands. Terminalia. Tergal lobes of IX each
with approximately 13-17 setae; basimere with a slightly-defined basal tergo-
mesal lobe, bearing a number of slender setae; claspette filament not notice-
ably wider at any particular point, tapering to apex from before middle.
PUPA. No specimens available.
LARVA. Not seen. Brug (1931) describes and figures the single lar-
val skin associated with an adult of nigrorhynchus as follows: Head. Hairs
d, 6-C with 6 branches; 14-C stellate, with 4; basal maxillary hair stellate,
with 8. Thorax. Hair 1-P with 3 branches; 3-P with 2. Abdomen. Comb
scales pointed, with a short lateral fringe; pecten with last tooth simple and
no teeth inserted beyond siphon hair tuft; siphon hair tuft reaching or exceed-
ing siphon apex; saddle with 4 spines laterally on distal margin; hair 1-X con-
siderably longer than saddle, double. Body pilosity is not mentioned or fig-
ured.
TYPE DATA. Type is non-existent; type locality: Djajasana,
Preanger Regentschappen, 1,400 meters (Thienemann), JAVA.
DISTRIBUTION. Specimens examined: JAVA, Djajasana and
Tangkoekan Prahoe, 3 males, 5 females. Records from literature: JAVA
(Brug 1931).
TAXONOMIC DISCUSSION. Based on the discovery that both the
males and females of nigrorhynchus differ from harveyi in having the prealar
and upper sternopleural scale patches distinctly separated, I have here given
this form the status of a full species.
Since associated larval skins have not been seen for both sexes of this
Species, it is still conceivable that the male and female described here as
representing nigrorhynchus are actually not conspecific. A fuller understand-
ing of this species will not be possible until series of adult-associated larval
specimens are available.
BIOLOGY. The type collection originated from larvae collected from
water in a tree hole.
Za Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
AEDES (FINLAYA) RIZALI (BANKS)
(Figures 2, 9)
Culex rizali Banks 1906, Philipp. J. Sci. 1: 999 (¢).
Aedes (Finlaya) abadsantosi Baisas 1946, Mon. Bull. Philipp. Hlth. Serv.
22(3): 25 (o, 2, P*, L*); Knight 1947 (1948), Ann. ent. Soc. Amer. 40:
641 (o*, 9, L*), NEW SYNONYMY.
Aedes (Finlaya) burgosi Baisas 1946, Mon. Bull. Philipp. Hlth. Serv. 22(3):
27 (o', 2, P, L*); Knight and Hull 1951, Pacif. Sci. 5: 240 (<, L*).
NEW SYNONYMY.
FEMALE. Head. Torus with dark scales mesally; palpus with a few
pale scales apically; proboscis dark scaled; vertex with upright forked scales
usually dark. Thovax. Anterior pronotum with a few narrowed yellowish
scales above, broad creamy scales below; posterior pronotum dorsally and
posteriorly with narrow yellowish scales, some of posterior scales may be
broadened, occasionally a few dark scales medially; median scutal line ex-
tending across prescutellar area to near scutal border; submedian line
broken at level of scutal angle, posterior portion bent antero-laterally along
scutal angle to a variable extent, sometimes connecting anteriorly with
lateral line; prealar scale patch distinctly separated from upper sternopleu-
ral scale patch (Figure 2); paratergite and subspiracular area not scaled.
Legs. Fore and mid femora anteriorly dark, occasionally a few scattered
pale scales present; hind femur with apico-ventral pale-scaled area not
broadly expanded onto anterior aspect; fore and mid tarsi with narrow basal
white rings on I-II, sometimes a few pale scales ventrally at base of III of
mid legs; hind tarsus with broad basal bands on I-III, that on III occupying
basal 0.4 of segment (partially broken ventrally by dark scales). Abdomen.
Tergum I dark dorsally, II usually dark but sometimes with basal pale scal-
ing; IJI- VIII with complete basal white bands, occasionally bands somewhat
narrowed medially, and more occasionally even incomplete on 1 or more
segments. Variation. In the British Museum collection there is a female
specimen from the Philippines that resembles vizalz except that the anterior
portion of the postspiracular scale patch consists of narrow yellowish scales
and does not extend to the ventral margin of the spiracle. All of the abdomi-
nal terga are dorsally dark except for a complete basal band on VIII. This
specimen has the following data: Mt. Mupo, Dansalan (Mindanao) 28. III. 1920
(Dr. A. Moore). In the collection of the Academy of Natural Sciences of
Philadelphia, there is a female specimen, Lagolago, Leyte (H. R. Roberts),
with the postspiracular scales similar to those of the above specimen but
differing in having the terga with complete basal bands.
MALE. Head. Palpus dark with white scales over joint of II-III, a
latero- and ventro-basal white-scaled area on IV; proboscis dark scaled.
Abdomen. Tergum I dark, II with medially incomplete basal band, IT-VIII
with complete basal bands. Terminalia. (Figure 9.) Tergal lobes of IX
each with approximately 5-10 setae, but it should be noted that 3 of the type
series of buvgosi were described as having ''more than 20-30 bristles" on
each lobe. Claspette filament slightly enlarged just beyond middle.
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 23
Variation. In the USNM there is a male specimen, with associated larval
and pupal skins, which cannot be separated from vizali except on male geni-
talic characters. The claspette blade is slender and tubular instead of being
laterally flattened and there are 11-13 hairs on each of the tergal lobes of IX
(San Ramon, Mindanao, 17. IX. 45).
PUPA. Chaetotaxy similar to saxicola (Figure 11).
LARVA. (Figure 9.) Body clothed with stellate hairs and pile (vari-
ably developed). Head. Hairs 5,6-C with 7-8 branches; 14-C and basal
maxillary hair (bmh) stellate, with 4-7. Thorax. Integument heavily pilose,
pile simple or branched; hair 1-P with 4-5 branches; 3-P with 5-8; 8-P stel-
late, with 7-11; 14-P stellate, with 3-7; 1-M stellate, with 6-10; 14-M stel-
late; 1-T stellate, with 10-13; 5-T stellate, with 5-9; 7-T with 5-9, similar
in shape and development to 6-M. Abdomen. Integument heavily pilose, pile
simple or branched; many stellate hairs; 1-III-VII with 2-5 branches; comb
consisting of 50-72 scales, each scale broad, not appreciably tapered apical-
ly, with a conspicuous lateral and apical fringe, a medial heavier spicule
sometimes apparent; siphon pilose, index approximately 2. 7-3. 2; siphon
hair tuft with tips exceeding siphon apex; pecten with 15-20 teeth, 4-6 teeth
beyond siphon hair tuft, apical 4-7 teeth simple, remainder each with 1-3
ventral denticles (may be hard to see unless siphon is strongly flattened),
apical 2-5 teeth ventrally out of line with others, tip of apical tooth approxi-
mately on level with siphon apex; hair 1-X with 2 equal branches, rarely 3,
longer than saddle; ventral brush without precratal tufts. Variation. In the
type description of burgosi, Baisas included a small series of specimens
from Mindanao (but not from the type locality). Two whole larvae from this
series are inthe USNM. These 2 larvae differ from one another on the basis
of the shape of the comb teeth. One (R112-k; figured by Knight and Hull 1951:
241) has comb teeth with a stout central spicule, whereas the comb teeth of
the other (R112-m) have a more even apical fringe. The two also differ from
one another quite distinctly on the numbers of branches of various hairs.
Together, these two specimens differ from both vizali and sherki because the
apical pecten teeth are in line, or if anything, slightly dorsally removed (not
ventrally out of line). In view of the large numbers of specimens of vizali
and sherki seen by me and of the fact that none of them deviated from having
the apical teeth ventrally out of line, it would seem that this could be a dif-
ference at the species level. However, present information is not sufficient
to permit the reconciliation of these discrepancies.
TYPE DATA. Culex rizali Banks, type non-existent; type locality:
Volcano Canlaon, Mt. Siya-Siya, at altitude of 760 m, Negros Occidental
Province, Negros Island (Banks), PHILIPPINES. Aedes abadsantosi Baisas,
type non-existent; type locality: Llavac, Infanta Muni. , Tayabas Province,
Luzon Island (Baisas), PHILIPPINES. Aedes burgosi Baisas, type non-ex-
istent; type locality: Titunod Creek in Kolambugan, Lanao, Mindanao Island
(Guinto), PHILIPPINES.
DISTRIBUTION. Specimens examined: PHILIPPINES, Samar,
Mindanao, Leyte, 26 males, 70 females, 25 larvae, 40 rearings. Records
from literature: PHILIPPINES, Negros (Banks 1906), Luzon and Mindanao
(Baisas 1946).
24 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
TAXONOMIC DISCUSSION. The absolute identity of this species
remains in doubt because the 2 female specimens upon which it was founded
were destroyed during the Second World War liberation of Manila and no
topotypic material has yet become available. However, Banks' description
was unusually precise (for that period) and it is possible to recognize rather
conclusively that he was dealing with the widely distributed Philippine spe-
cies described above. Some of the Palawan specimens designated as "novo-
types'' for this species by Baisas (1946) were examined by Knight (1947) and
found to be conspecific with saxicola.
A. abadsantosi was described by Baisas as being distinct from vizali
in having the median scutal golden line thickened but not forked at the pre-
scutellar area, the submedian and lateral scutal lines not meeting anteriorly
to form a loop, and abdominal tergum VIII of the female always possessing a
broad white basal band. Regarding the first character, Banks does not de-
scribe the median scutal golden line as being forked posteriorly, but states
as follows: "A narrow, golden median line from anterior margin halfway
through posterior bare brown spot. . ."' In connection with the submedian
and lateral scutal lines forming an anterior loop in vizali, the extent to which
the submedian line breaks and bends laterally at the scutal angle varies wide-
ly in the large series of Philippine specimens seen by me and there is no in-
dication that it is other than a rather variable character.
This is also true of the extent of the basal white markings on the fe-
male abdominal terga. Banks described vizali as having the abdominal terga
dark except for baso-lateral white patches on VI-VIII. A. abadsantosi is
described as having tergum VIII constantly with a broad white basal band, V-
Vil sometimes also banded basally, and III-IV with scattered basal pale
scales. Fifty-six females with abdomens suitable for examination were
available to me. Among these specimens dorsal abdominal markings varied
from complete basal banding on segments II-VIII to having only a narrow
band on VII. The extent of dorsal abdominal markings is a notoriously vari-
able character at best, and no evidence presently exists that this character
can be used here for separation of species.
Although it appears that nearly the entire range of variation can be
found in any single population, it is true that northern populations (Luzon,
Leyte, and Samar) have a high percentage of individuals with complete band-
ing on at least segments II-VI, and southern populations (Negros and
Mindanao) have fewer segments banded and such bands or markings as occur
are less well developed. It is possible that incipient species are developing
and even now are biologically distinct. However, in the absence of biological
studies of the variation found here, it is considered necessary to treat all of
this material as a single species.
A .burgosi was considered to be distinct from vizali and abadsantosi
by Baisas on the basis of having more bristles on the ninth tergum. However,
here again a perfect continuum exists and burgosi is accordingly synonymized
to vizalt.
BIOLOGY. The larva is commonly found in the water collected in
rock holes in and along the stream beds of forested streams. The rock pools
may be small or large, and in shade or sun. Occasionally, the larvae are
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 25
also found in hollows on roots or fallen logs along stream beds.
Nothing has been reported on the habits of the adults except that the
2 females of the type series of vizali were collected in the act of biting
(Banks 1906).
AEDES (FINLAYA) SAXICOLA EDWARDS
(Figures 10, 11)
Hulecoeteomyia fluviatilis Leicester (non Lutz 1904) 1908, Cul. Malaya: 111
io 9),
Aedes (Finlaya) saxicola Edwards 1922, Indian J. med. Res. 10: 262 (nomen
novum for fluviatilis); Barraud 1934, Fauna Brit. India 5: 191 (o*, ?*,
L*); Knight 1947 (1948), Ann. ent. Soc. Amer. 40: 628 (o*, 2, L*);
Knight and Hull 1951, Pacif. Sci. 5: 239 (taxonomy).
Finlaya greigit Barraud 1923, Bull. ent. Res. 13: 406 (<, ?); Edwards 1932,
in Wytsman, Genera Insect. fasc. 194: 151 (synonymy).
Aedes (Finlaya) rizali Banks, Baisas 1946, Mon. Bull. Philipp. Hlth. Serv.
22(3): 21 (misidentification).
FEMALE. Head. Torus with dark scales mesally; palpus with a few
pale scales apically; proboscis all dark scaled; vertex with medial upright
forked scales usually dark. Thorax. Anterior pronotum with broad white
scales; posterior pronotal scaling extremely variable, ranging from narrow
to narrow-curved and even to broad scales, these scales may all be yellow-
ish or all brownish or with all combinations in between, they may be numer-
ous over entire dorsal portion of area or scanty; median scutal line forked
at prescutellar space, sometimes appearing posteriorly tapered; submedian
line broken at level of scutal angle with anterior end of its posterior portion
bent briefly antero-laterally along scutal angle, anterior portion more weak-
ly developed than posterior portion, sometimes nearly obsolescent; line over
wing base usually rather diffused, sometimes even obsolescent, anteriorly
well separated in area of scutal angle from submedian line; prealar scale
patch distinctly separate from upper sternopleural scale patch; paratergite
and subspiracular area without scales. Legs. Fore femur anteriorly dark,
with a baso-ventral line of pale scaling; mid femur usually with some pale
scales along anterior midline; hind femur with anterior submedial white area
sometimes connecting dorsally with posterior white area, apico-ventral pale
area extensively produced onto anterior surface; fore and mid tarsi with dis-
tinct basal white areas on I-II; hind tarsus with prominent basal white bands
on I-III, that on III dorsally occupying approximately half or more of the seg-
ment, a few basal white scales may be present onIV. Abdomen. Terga
dorsally dark scaled, baso-lateral silvery-white spots usually visible dor-
sally on VII-VIII, those on VIII sometimes forming a complete band; often a
small medio-basal patch of brownish-yellow scales on some terga. Vavria-
tion. A female specimen (00474-100) in the USNM from Thailand perfectly
resembles this species except the prealar and upper sternopleural scale
patches are continuous without any evidence of discontinuity. The anterior
26 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
surface of the mid femur and the dorsal aspect of the terga are entirely dark
scaled. There is a similar female specimen (33) in the BM from Hainan
Island. This differs from the first, however, in having a medio-basal patch
of brownish-yellow scales dorsally on terga III-VII. Since the confluence or
discontinuity of the prealar and upper sternopleural scale patches is seldom
subject to variation within the chrysolineatus subgroup, it is possible that
these specimens represent a presently unrecognized species. A series of
specimens from the Thailand changwats of Chanthaburi and Nakhon Si
Thammarat include in their number several individuals which have 2 sub-
spiracular scales, tergal basal bands (laterally disconnected from baso-
lateral spots), and broad dark scales dorso-posteriorly on the anterior pro-
notum. Some of the males of this series have ventral pale scaling along both
IV and V.
MALE. Head, Palpus dark, baso-ventral white scaling on terminal
segment; proboscis dark scaled. Abdomen. Terga dorsally dark scaled,
with a baso-medial spot of pale scales on II-VI, may be reduced to just a
few pale scales on some segments, baso-medial spots may occasionally form
narrow basal white bands by connecting laterally with baso-lateral spots.
Terminalia, (Figure 10.) Tergal lobes of IX each with approximately 7-14
setae; claspette filament with taper beginning at or just beyond middle.
Variation. ‘Two male specimens from Malaya have the anterior surface of
the middle femur all dark.
PUPA. Chaetotaxy as shown in Figure 11.
LARVA. (Figure 10.) Head. Integument with a noticeable sane
like rugosity; hairs 5, 6-C with 5-8 branches; 14-C and basal maxillary hair
(bmh) well developed, somewhat stellate in form, 14-C with 2-5 branches,
bmh with 3-7. Thovax. Without obvious pilosity; hair 1-P with 3-4 branches;
3-P with 4-5; 8-P with 2-4; 14-P with 2-4; 1-M with 2-4 elongate very stout
barbed branches; 1-T quite variable in development, with 1-6; 5-T with 1-2;
7-T very stout, about half length of 6-M, with 3-5 heavily barbed rather
blunt-ended branches, very unlike 6-M in shape and development. Abdomen.
Integument with fine unbranched pile, increasing posteriorly; hair 1-I-II
rather stellate, elongate on HI-VU, with 1-4 branches; 13-I-II short, some-
what stellate; comb consisting of 53-70 scales, each scale narrow, elongate,
slightly tapered, with a lateral and apical fringe, sometimes with a short
stout apical spicule; siphon non-pilose, index approximately 2. 5-3. 3; siphon
hair tuft with tips not attaining siphon apex; pecten with 14-19 teeth, 3-6
inserted beyond attachment of siphon hair tuft (are the largest and may be ir-
regularly spaced), more basal teeth with 1-2 very small ventral denticles
medially; hair 1-X stout, barbed, considerably longer than saddle, single;
ventral brush without precratal tufts. Variation. Very extensive variation
occurs in the development of thoracic and abdominal hairs. One form has
1-T, hair 1-IJI-VII, and hair 6-I-VI all strongly developed and single; and
other hairs in general showing a reduction in numbers of branches. Con-
versely, a true stellate form exists in which the following hairs become elon-
gate, stellate, and possessed of many more branches: 8-P; 14-P; 1-M;
14-M; 1-T; 13-T; 1,2,5, 11, 13-I-I]; and 2,5-IN-VI. Fairly conspicuous body
pilosity can also occur on these larvae. The development of "hairy" forms
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 27
of mosquito larvae has been discussed by Rosen and Rozeboom (1954),
Colless (1956), and Reid (1963), and it is apparently an environmentally-in-
duced condition.
TYPE DATA. Hulecoeteomyia fluviatilis Leicester, type non-exis-
tent; type locality: Ulu Gombak (Leicester), MALAYSIA. Finlaya greigi
Barraud, male and female cotypes in British Museum; type locality: Haflong,
Cachar Hills, Assam (Barraud), INDIA.
DISTRIBUTION. Specimens examined: PHILIPPINES, Palawan, 7
males, 9 females, 7 larvae, 5 rearings. MALAYSIA, Selangor, Cameron
Highlands, 4 males, 3 females, 3 rearings. THAILAND, Chumphon, Banong,
Narathiwat, Pattalung, Songkhla, Chiang Mai, Nakhon Nayok, Trad, Ranong,
Pathum Thani, Trang, Khaulan, Chanthaburi, Nakhon Si Thammarat, 19
males, 77 females, 168 larvae, 38 rearings. JAVA, Tjisanoea, Ngadisari,
2 males, 3 females. BORNEO, Lg. Tebangan, Sarawak and Heningau, N.
Borneo, 6 males, 9 females. SUMATRA, Arastagi, 1 male. Records from
literature: INDIA, Eastern Himalayas, Assam (Barraud 1934). MALAYSIA,
Wray's Hill, Pahang (Edwards 1928). SINGAPORE, Pulai Oban (Edwards and
Given 1928). JAVA, Mt. Salak (Barraud 1934). THAILAND (Causey 1937).
TAXONOMIC DISCUSSION. This species, while nearest to A.
jugraensis, more or less stands alone in the chrysolineatus subgroup. Al-
though subject to extensive variation, the adult is distinctive in having the
proboscis all dark scaled, the median longitudinal line forked posteriorly,
and the mid femur with the anterior and antero-dorsal surfaces dark apically.
The larva is distinctive from all of the others except jugvaensis in the highly
modified form and development of hair 7-T.
BIOLOGY. The larva is commonly found in rock holes and pools in
the beds of jungle streams. These pools may be very small or large, may
be entirely in shade or at least part of the time in the sun, and may be with
or without leaves on the bottom. On occasion, it may occur in tree holes
and in hollows on top of fallen logs. One or 2 records each exist for such
different places as coconut half shells, a water jug, bamboo, seepage in
small hole, and ina sump. Whether these represent errors or valid collec-
tions cannot now be determined. The adult female has been taken on several
occasions in human biting collections in forest.
AEDES (FINLAYA) SHERKI KNIGHT
(Figures 7, 12)
Aedes (Finlaya) sherki Knight 1947 (1948), Ann. ent. Soc. Amer. 40: 645
(oe; Lae
FEMALE. Head. Torus with dark scales mesally; palpus with a few
pale scales apically, occasionally a few pale scales apically on penultimate
segment; proboscis all dark scaled; upright forked scales of vertex usually
dark. Thorax. Anterior pronotum with a few narrowed yellowish scales
above, broad creamy scales below; posterior pronotum sparsely covered
with narrow-curved yellowish scales, some of posterior scales may be
28 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
broadened, occasionally a few dark scales medially; median scutal line
extends posteriorly nearly across prescutellar area, not forked posteriorly;
submedian line broken at level of scutal angle with its posterior portion bent
briefly antero-laterally along the angle, line over wing base anteriorly does
not connect at scutal angle with outcurved portion of submedian line; prealar
scale patch distinctly separate from upper sternopleural scale patch; para-
tergite and subspiracular area without scales. Legs. Fore femur anteriorly
dark, with a baso-ventral line of pale scaling; mid femur anteriorly dark;
hind femur with apico-ventral pale area broadly produced anteriorly but not
ringing apex; fore tarsus and sometimes mid tarsus with a small basal patch
of white scales on III (as well as on I-II); hind tarsus with first 4 tarsomeres
basally banded. Abdomen. Terga dorsally dark scaled, sometimes with a
small medio-basal creamy patch on 1 or more of segments, VII with baso-
lateral silvery-white spots extending prominently onto dorsum; VIII witha
complete basal white band.
MALE. Head. Palpus dark, a few pale scales over joints between
II-III and III-IV; proboscis dark scaled. Abdomen. Terga with complete
broad basal white bands on III-VI, sometimes bands narrowed dorsally or
with dorsal portion separated from dorso-lateral spot on 1 or more segments.
Terminalia. (Figure 7.) Tergal lobes of IX each with approximately 9-14
setae; claspette filament slightly widened medially.
PUPA. Chaetotaxy similar to saxicola (Figure 11).
LARVA. (Figure 12.) Body clothed with stellate hairs and with pile.
Head, Hairs 5, 6-C with 5-8 branches; 11-C, 14-C, and basal maxillary hair
(bmh) stellate, 14-C with 3-6 branches, bmh with 6-10. Thorax. Integument
heavily pilose, pile unbranched; hair 1-P with 4-9 branches; 3-P with 5-13;
8-P stellate, with 6-12; 14-P stellate, with 5-11; 1-M stellate, with 2-12;
1-T stellate, with 9-16; 5-T stellate, with 6-11; 7-T with 7-16, similar in
shape and development to 6-M. Abdomen. Integument heavily pilose, pile
simple or branched, many stellate hairs; 1-I-III stellate; 5-IV-VII elongate,
with 5-7 branches on IV, 3-5 on V, 3-7 on VI-VIJ; comb consisting of 62-73
scales, each scale narrow, elongate, with a conspicuous lateral fringe, api-
cal spicule sometimes conspicuously more developed than adjacent lateral
spicules; siphon pilose, index 3. 4-4. 0; siphon hair tuft with tips extending
beyond siphon apex; pecten with 15-20 teeth, 4-5 teeth beyond siphon hair
tuft, most of teeth with 1-3 ventral denticles, apical 2-3 teeth ventrally out
of line with others, tip of most apical tooth usually extending beyond siphon
apex; hair 1-X with 2, rarely 3, equal branches, longer than saddle; ventral
brush without precratal tufts.
TYPE DATA. Holotype male in U. S. National Museum; type locality:
Baguio, City of Baguio Province, Luzon Island (Rozeboom), PHILIPPINES.
DISTRIBUTION. Specimens examined: PHILIPPINES, Luzon, 51
males, 57 females, 17 larvae, 5 rearings.
TAXONOMIC DISCUSSION. This species is very closely related to
rizali, differing from that species only in possessing basal white bands on
hind tarsomeres I-IV (instead of only on I-III).
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 29
BIOLOGY. As with vizali, the larva is found in the water collected
in rock holes in and along the stream beds of forested streams. Records
also exist of the larva occurring in artificial containers.
ACKNOWLEDGEMENTS
Iam grateful to Dr. Botha de Meillon, Responsible Investigator,
South East Asia Mosquito Project and to Dr. Alan Stone, Agricultural
Research Service, U. 8. Department of Agriculture, for their many helpful
suggestions and criticisms in connection with this paper. The loan of im-
_ portant collections from Indonesia and Borneo by Dr. P. F. Mattingly,
Department of Entomology, British Museum (Natural History) is acknowl-
edged with sincere appreciation. The examination of types in Tokyo by Dr.
Mercedes D. Delfinado and in the British Museum (Natural History) by Dr.
Ralph A. Bram, both of the South East Asia Mosquito Project, was of great
value to this study. The illustrations were prepared by Mrs. Jung Lea
Smith and Mr. Vichai Malikul of the South East Asia Mosquito Project. Miss
Helle Starcke typed the manuscript for offset reproduction.
LITERATURE CITED
BAISAS, F. E.
1946. Notes on Philippine mosquitoes, X. Some species of Aedes
(Finlaya) and (Aedimorphus). Mon. Bull. Philipp. Hlth. Serv.
22(3): 21-37.
BANKS, C. S.
1906. A list of Philippine Culicidae with description of some new
species. Philipp. J. Sci. 1: 999-1003.
BARRAUD, P. J.
1923. A revision of the culicine mosquitoes of India. Part VIL.
Indian J. med. Res. 11: 495-505.
1924. A revision of the culicine mosquitoes of India. Part VIII.
Indian J. med. Res. 11: 845-865.
1934. The fauna of British India including Ceylon and Burma.
Diptera 5, family Culicidae, tribes Megarhinini and Culicini.
Taylor and Francis, London, 463 pp.
BELKIN, J. N.
1962. The mosquitoes of the South Pacific. Univ. Calif. Press,
Berkeley, 2 vols. , 608 and 412 pp.
BOHART, R. M.
1959. A survey of the mosquitoes of the Southern Ryukyus. Mosquito
News 19: 194-197.
BOREL, E.
1928. Les moustiques de la Cochinchine et du Sud-Annam. — II.
Aedes (Finlaya). Arch. Inst. Pasteur Indo-Chine 8: 41-76.
1930. Les moustiques de la Cochinchine et du Sud-Annam. Monogr.
Coll. Soc. Pat. exot. 3, 423 pp., Paris.
30 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
BRUG, S. L. :
1931. XXXII. Culiciden der Deutschen Limnologischen Sunda-
Expedition. Arch. Hydrob. 9: 1-42.
BRUG, S. L. and F. W. EDWARDS.
1931. Fauna Sumatrensis. Culicidae (Diptera). Tijdschr. Ent. 74:
241-261.
BRUG, S. L. and J. BONNE-WEPSTER.
1947. The geographical distribution of the mosquitoes of the Malay
Archipelago. Chron. nat. 103: 179-197.
CARTER, H. F. and D. P. WIJESUNDARA.
1948. Notes on some Ceylon culicine mosquitoes. Ceylon J. Sci.
(B) 23: 135-151.
CHAGIN, K. P. and P. I. KONDRATIEV.
1943. Vectors of autumnal (Japanese) encephalitis in Soviet Far
East and their control. Med. Parasit. and Parasitic Dis. 12:
34-44, Abstract in Biological Abstracts 19(8), No. 17661,
Oct. 1945.
CAUSEY, O. R.
1937. Some anopheline and culicine mosquitoes of Siam with re-
marks on malaria control in Bangkok. Amer. J. Hyg. 20:
400-420.
CHOW, C. Y. ,
1949. Culicine mosquitoes collected in Western Yunnan, China dur-
ing 1940-1942. Proc. ent. Soc. Wash. 51: 127-132.
1950. Collection of culicine mosquitoes (Diptera, Culicidae) in
Taiwan (Formosa), China, with description of a new species.
Quart. J. Taiwan Mus. 3: 281-287.
COLLESS, D. H.
1956. Environmental factors affecting hairiness in mosquito larvae.
Nature 177: 229-230.
EDWARDS, F. W.
1928. Mosquito notes. — VII. Bull. ent. Res. 18: 267-284.
EDWARDS, F. W. andD. H. C. GIVEN.
1928. The early stages of some Singapore mosquitos. Bull. ent.
Res. 18: 337-357.
EDWARDS, F. W.
1932. Genera Insect. Diptera, Fam. Culicidae. Fascicle 194.
Desmet-Verteneuil, Brussels. 298 pp.
FENG, L-C.
1938. The tree hole species of mosquitoes of Peiping, China. Chin.
med. J. 2: 503-520.
KNIGHT, K. L. |
1947 (1948). The Aedes (Finlaya) chrysolineatus group of mosquitoes
(Diptera: Culicidae). Ann. ent. Soc. Amer. 40: 624-649.
Knight: Finlaya Subgroup chrysolineatus of Southeast Asia ot
KNIGHT, K. L. and W. B. HULL.
1951. The Aedes mosquitoes of the Philippine Islands. I. Keys to
species. Subgenera Mucidus, Ochlerotatus, and Finlaya
(Diptera, Culicidae). Pacif. Sci. 5: 211-251.
KNIGHT, K. L. and E. N. MARKS.
1952. An annotated checklist of the mosquitoes of the subgenus
Finlaya, genus Aedes. Proc. U. S. nat. Mus. 101: 513-574.
LACASSE, W. J. and S. YAMAGUTI.
1950. Mosquito fauna of Japan and Korea. Off. Surgeon, 8th U. S.
Army, Kyoto, Honshu. 268 pp.
LEICESTER, G. F.
1908. The Culicidae of Malaya. Stud. Inst. med. Res. F.M.S. 3:
18-261.
LIEN, J. C.
1962. Non-anopheline mosquitoes of Taiwan: annotated catalog and
bibliography. Pacif. Ins. 4: 615-649.
MACDONALD, W. W.
1957. Malaysian parasites — XVI. An interim review of the non-
anopheline mosquitoes of Malaya. Stud. Inst. med. Res. F.
M.S.,No. 28, pp. 1-34.
MACDONALD, W. W. andR. TRAUB.
1960. Malaysian parasites — XXXVII. An introduction to the ecol-
ogy of the mosquitoes of the lowland dipterocarp forest of
Selangor, Malaya. Stud. Inst. med. Res. F.M.S., No. 29,
pp. 79-109.
REID, J. A.
1963. Notes on anopheline mosquitoes from Malaya, with descrip-
tions of three new species. Ann. trop. Med. Parasit. 57:
97-116.
ROSEN, L. and L. E. ROZEBOOM.
1954. Morphologic variations of larvae of the scutellaris group of
Aedes (Diptera, Culicidae) in Polynesia. Amer. J. trop.
Med. and Hyg. 3: 529-538.
THEOBALD, F. V.
1911. Culicidae of the Royal Zoological Society ''Natura Artis
Magistra'' Amsterdam and description of three new species.
Tijdschr. Ent. 54: 233-240.
TSAI, C. and J. LIEN.
1950. A new species of Aedes (Finlaya) found in Taiwan. J. med.
Ass. Formosa 49: 177-188.
YAMADA, S.
1921. Descriptions of ten new species of Aedes found in Japan, with
notes on the relation between some of these mosquitoes and
the larva of Filaria bancrofti Cobbold. Annot. zool. jap. 10:
45-81.
32 Contrib. Amer. Ent. Inst., vol. 2, no. 5, 1968
YAMADA, §.
1927. An experimental study on twenty-four species of Japanese
mosquitoes regarding their suitability as intermediate hosts
for Filaria bancrofti Cobbold. Sci. Rep. Inst. infect. Dis.
6:559-622.
formosensis
chrysolineatus
/Enens/s
e
hint
japonicus §
Jugraensis
Fig. 2
jugraensis
1S
tienensis
formosens
Ain
d femur)
japonicus s
(mi
lineatus
chryso
ind femur)
(h
jugraensis
enensis
ti
Ain
(hind femur)
japonicus s
d femur)
°
in
(h
E
E
°
O.Imm
chrysolineatus
10
VA 12
10
12
ss
£0
VMauKu_—
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UL. SMITH
mayne
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y
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Vill
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i
a BASMITA
LEER ag RS
o = es ee SSS
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re
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Knight: Finlaya Subgroup chrysolineatus of Southeast Asia 45
INDEX
Valid names are set in roman type, synonyms are italicized.
The italicized pages are those which begin the primary treatment of the
Species. Numbers in parentheses refer to the figures illustrating the
Species in question.
abadsantosi 22,23, 24
aureostriatus subgroup 2
burgost 22,23, 24
— chrysolineata 8,9
chrysolineatus 4,5, 6, 8,9, 10, 12, 14, 15, (1, 2, 3)
chrysolineatus subgroup 2, 5, 10, 20, 26, 27
Colocasia 10,12, 13
Culex 8,9, 15, 22, 23
eucleptes 15, 16
fluviatilis 25, 27
formosaensis 10
for mosensis 5,8, 1, 10, 48. 18.15, 4,2, 4)
gvreigi 25,27
harveyi 5, 6, 8, 10,22, 14, 15,21, (5)
harveyi var. nigrorhynchus 20
Howardina 8,9
Hulecoeteomyia 8, 9, 19) 25, 27
japonicus 15, 16,17
japonicus japonicus 5, 6, 7, 15, 18, (6)
japonicus shintienensis 5,8, 2741. a.
japonicus var. ceylonica 8,9
jugraensis 6, 7,29, 21, 1,.2,8)
Rhasiana 10, 12
koreicus | 0, 4,17
Musa ip
nigrorhynchus 9, 6, 7,20, 21
pallirostris 10, 12
rizali 6.7, 19, 22,23) 26, 25, 26) 29, (2.9)
saxicola 5, 6, 7,9, 11, 13, 20, 23, 24, 25, 28, (10, 11)
sherki Db aah erly te)
shintienensis 7
togoi subgroup 2
trilineata 8,9
Wuchereria bancrofti 17
MAILING DATES FOR CONTRIBUTIONS, VOLUME 1
No. 1. Oct. 7, 1964 No. 5. Sept. 9, 1965
No. 2. June 21, 1965 No. 6. Aug. 12, 1966
No. 2a. June 6, 1967 No. 7. March 28, 1967
No. 3. Sept. 15, 1965 No. 8. June 5, 1967
No. 4. June 25, 1965
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