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Discovering Diverse Content Through
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Scales.—The most important structural peculiarities in Culicidæ are the scales, which
form the chief and most readily observed characters for separating genera and species.
The importance of scale structure has been recently ignored by some workers, who are
probably right academically, but as a means of separating groups, and so more easily
running down a species, the practical man is strongly advised to follow this method. As
to what a genus is, is purely a matter of personal opinion. If one examines any recent
standard work on entomology one will find a species being placed in varied genera by
the varied authorities.
The head, thorax, abdomen and wings are in nearly all cases clothed with squamæ of
varied form, of which the following are the main types (fig. 397):—
(1) Flat, spade-shaped scales (a).
(2) Narrow curved scales (e).
(3) Hair-like curved scales (d).
(4) Spindle-shaped scales (f).
(5) Small spindle-shaped scales (g).
(6) Upright forked scales (h) and (i).
(7) Twisted upright scales (j).
(8) Inflated or pyriform scales (k).
(9) Mansonia scales (b).
(10) Small broad asymmetrical scales (c).
Various other varieties are found on the wings, such as:—
(1) Narrow linear lateral scales.
(2) Narrow lanceolate scales.
(3) Broad lanceolate scales.
(4) Elongated, broad, truncated scales (= Pseudotæniorhynchus-like scales).
(5) Pyriform scales.
(6) Asymmetrical broad or Tæniorhynchus scales.
(7) Flat spade-like scales. 389
The wings have a series of scales along the middle line of the veins, and also lateral
scales to all or nearly all the veins. The wing is also fringed by a series of scales
(fig. 396), which, however, are of little systematic importance; the so-called “border
scales” (b.s.) vary, however, to some extent, and are useful characters in separating
some of the Tæniorhynchus.

The Classification of Culicidæ.

Section A.—Proboscis formed for piercing; metanotum nude. Scutellum simple.
I. Wings with six-scaled longitudinal veins.
A. Palpi long in the male.
α. Palpi long in both sexes, clavate in ♂ Anophelina.
I. First submarginal cell as long or longer than the second posterior cell.
Antennal segments without dense lateral scale tufts.
Wing scales
lanceolate Anopheles, Meigen.
Wing scales mostly
long and narrow Myzomyia, Blanchard.
Prothoracic Wing scales as
lobes above, but fourth
Thorax and
simple; no long vein near
ab­do­men
flat head base of third and
with
scales outstanding scales
hair-like
on prothoracic
scales
lobe Neomyzomyia, Theobald.
Wing scales partly
large and inflated Cycloleppteron, Theobald.
Prothoracic lobes mammillated;
some flat head scales. Basal lobe
of ♂ genitalia of two segments Stethomyia, Theobald.
Prothoracic lobes with dense outstanding scales Feltinella, Theobald.
Thorax with some narrow curved scales;
abdomen hairy Pyretophorus, Blanchard.
Wing scales small and lanceolate. Wing scales
broad and lanceolate Myzorhynchella, Theobald.
Thorax with hair-like curved scales, some narrow
curved ones in front; abdomen with apical
lateral scale tufts, scaly venter; no ventral tuft Arribalzagia, Theobald.
Thorax with hair-like curved scales; abdominal
scales on venter only, with a distinct ventral
apical tuft Myzorhynchus, Blanchard.
Much as above, but abdomen with long spine-like
dense lateral tufts Chrystia, Theobald.
Thorax with very long hair-like curved scales;
abdomen pilose, except last two segments
which are scaly; dense scale tufts on third
femora; wings with broadish, blunt, lanceolate
scales Lophoscelomyia, Theobald.
Abdominal scales as lateral dorsal
patches of small flat scales;
Thorax and thoracic scales narrow and
abdomen curved, or spindle-shaped Nyssorhynchus, Blanchard.
with Abdomen nearly completely covered
 with Abdomen nearly completely covered
scales with irregular scales and with
lateral tufts Cellia, Theobald.
No lateral scale tufts Neocellia, Theobald.
Thoracic scales hair-like except a few narrow
curved ones in front; abdominal scales long,
broad and irregular Kerteszia, Theobald.
Thorax with hair-like curved scales and some
broad straight scales, others spatulate on
sides. Abdomen covered with fine hairs except
last three segments, which are scaly. Tufts of
scales on hind femora. Wing scales lanceolate Manguinhosia, Cruz.
Antennal segments with many dense scaly tufts Chagasia, Cruz.
Antennæ with outstanding scales on second
segment, more appressed ones on the first. At
least one segment of abdomen with long flat
more or less spatulate scales Calvertina, Ludlow.
II. First submarginal cell very small Bironella, Theobald.
With a distinct cylindrical tubercle projecting Dactylomyia, Newstead and
obliquely from the prothoracic region Carter. 390
Scutellum trilobed.
First submarginal cell much smaller than the
second posterior cell; proboscis long and bent Megarhininæ.
Palpi long in both sexes Megarhinus, Rob. Desvoidy.
Last segment of ♂ palpi blunt. Last segment of ♂
palpi long and pointed Ankylorhynchus, Lutz.
β. Palpi short in the female Toxorhynchites, Theobald.
First submarginal cell longer than the second
posterior cell Culicinæ.
Legs more or less densely scaly; head not
entirely clothed with flat scales; all the legs
densely scaly.
Wings with large pyriform scales Mucidus, Theobald.
Wings with narrow scales Psorophora, Rob. Desvoidy.
Hind legs only densely scaled Janthinosoma, Arribalzaga.
Head entirely clothed with flat scales. Legs
uniformly scaled with flat scales. Head and
scutellar scales all flat and broad. Palpi of ♀
short, of ♂ thickened apically and tufted Stegomyia, Theobald.
Palpi of ♀ longer than in Stegomyia and in ♂ long
and thin, acuminate, simple Desvoidea, Blanchard.
Head scales mostly flat, but a median line of
narrow curved ones; scutellar scales flat on
mid lobe, narrow curved on lateral lobes and
 palpi longer than proboscis Macleayia, Theobald.
Head scales mostly flat, irregular, narrow curved
ones behind; mid lobe scutellum with flat
scales, lateral with narrow curved; ♂ palpi
shorter than proboscis Catageiomyia, Theobald.
Head scales mostly flat, but a few narrow curved
ones in middle in front; scutellar scales all flat Scutomyia, Theobald.
Head scales all flat; scutellar scales all narrow
curved Skusea, Theobald.
Head with flat scales, except a small median area
of narrow curved ones; scutellar scales all
narrow curved Howardina, Theobald.
Head with all flat scales except a thin line of
narrow curved ones behind; scutellar scales all
narrow curved Danielsia, Theobald.
Head with small flat scales over most of surface,
with median line and line around eyes of
narrow curved ones; scutellar scales bluntly
spindle or club-shaped Hulecoetomyia, Theobald.
Head and scutellar scales narrow curved. Wing
scales long, narrowly lanceolate, collected in
spots; palpi clubbed in ♂ ; five-jointed and
rather long in ♀ Theobaldia, Neveu-Lemaire.
Wing scales (lateral) long and narrow, and ♀
palpi three-jointed, ♂ not clubbed and hairy Culex, Linnæus.
Wing scales at apex of veins dense and rather
broad, femora swollen; small dark species Melanoconion, Theobald.
Wings with short, thick, median scales and short,
broadish lateral ones on some of the veins;
scales mottled; fork-cells rather short Grabhamia, Theobald.
Wings with dense, broadish, elongated, truncated Pseudotæniorhynchus,
scales Theobald.
Wings with broad, short, asymmetrical scales Tæniorhynchus,
Arribalzaga.
Head covered with rather broad, flat, spindle-
shaped scales; scutellum with small flat scales
to mid lobe Gilesia, Theobald.
Head clothed with flat, irregularly disposed scales
all over, with patches of narrow curved ones; ♂
palpi clubbed Acartomyia, Theobald.
Abdomen with projecting flat lateral scales with
deeply dentate apices; wings not ornamented Lasioconops, Theobald.
Wings ornamented; scutellum with flat and
narrow curved scales Finlaya, Theobald.
γ. Palpi short in ♂ and ♀ Ædeomyina.
Wings unornamented.
Antennæ pilose in ♂ and ♀ ; second joint very
long Deinocerites, Theobald.
Antennæ plumose in the ♂.
Head clothed with narrow curved and flat scales.
Mid-lobe of scutellum with six border-bristles.
Scutellum with narrow curved scales.
Palpi in ♀ four-jointed, in ♂ two-jointed Ædes, Meigen.
Mid-lobe of scutellum with four border bristles.
Scutellum with flat scales.
Head clothed with flat scales only.
Fork-cells normal length.
Mid-lobe of scutellum with four border-bristles.
Palpi of ♀ two-jointed Verallina, Theobald.
Palpi of ♀ five-jointed, metallic Hæmagogus, Williston.
Fork cells very small or small.
Scutellar scales flat.
First submarginal cell longer than the second
posterior cell; no flat scales on mesothorax Ficalbia, Theobald.
First submarginal cell smaller than the second
posterior cell; flat scales on mesothorax Uranotænia, Arribalzaga.
Scutellar scales narrow curved.
First submarginal cell as in Uranotænia Mimomyia, Theobald.
Wings ornamented with Mansonia-like scales Ædeomyia, Theobald.
Section B.—Metanotum ornamented with chætæ, squamæ or both.
 α. With chætæ only.
Proboscis longer than whole body; lateral wing
scales Tæniorhynchus-like Phoniomyia, Theobald.
Proboscis as long as whole body in ♀ frons drawn
Binotia, Blanchard =
out into a prominence; wing scales rather
Runchiomyia, Theobald.
broad and long
Proboscis not as long as the whole body; lateral
vein scales narrow Wyeomyia, Theobald.
Proboscis not as long as whole body, swollen
apically; wing scales long and broad Dendriomyia, Theobald.
β. Metanotum with squamæ and chætæ.
Palpi short in ♂ and ♀.
Proboscis straight in ♀ and ♂ ; legs with scaly
paddles Sabethes, Rob. Desvoidy.
Venation like Sabethes.
Legs simple Sabethoides, Theobald.
Venation like Culex Goeldia, Theobald.
Proboscis in ♂ elbowed, with two scaly tufts Limatus, Theobald.
Palpi long in ♂, short in ♀ Joblotina, Blanchard.
Wings with seven-scaled longitudinal veins: Culex Heptaphlebomyia,
II.
type Theobald.

Section C.—Proboscis short, not formed for

piercing Corethrina.
Metatarsus longer than first tarsal joint Corethra, Linnæus.
Metatarsus shorter than first tarsal joint Mochlonyx, Ruthe. 391

Notes on the Different Genera.

Sub-family. Anophelina.
The following Anophelines have been recorded as malaria carriers:—
*Anopheles maculipennis, Meigen.
Anopheles bifurcatus, Linnæus.
*Myzomyia funesta, Giles.
Myzomyia lutzii, Theobald.
*Myzomyia rossii, Giles.
Myzomyia listonii, Liston.
Myzomyia culicifacies, Giles.
Pyretophorus superpictus, Grassi.
*Pyretophorus costalis, Loew.
Pyretophorus chaudoyei, Theobald.
 *Cellia argyrotarsis, Robineau Desvoidy.
Myzorhynchus pseudopictus, Grassi.
Myzorhynchus barbirostris, Van der Wulp.
Myzorhynchus sinensis, Wiedemann.
Myzorhynchus paludis, Theobald.
Myzorhynchus mauritianus, Grandpré.
Neocellia stephensii, Liston.
Neocellia willmori, James.
Nyssorhynchus theobaldii, Giles.
Nyssorhynchus fuliginosus, Giles.
Nyssorhynchus annulipes, Walker.
Those marked with an asterisk (*) also carry the larvæ of Filaria bancrofti, as also do
Myzorhynchus minutus, Theobald, and Myzorhynchus nigerrimus, Giles.

Genus. Anopheles, Meigen.

“Syst. Beschr. Europ. zwei. Ins. I,” 1818, ii, p. 2, Meigen; “Mono. Culicid.,” 1903,
i, p. 191; iii, p. 17; and 1910, v, p. 3, Theobald.
This genus contains a few large species found either in
temperate climates or in hills and mountains of warm
climates. The type is the European and North American
A. maculipennis.
Fig. 399.—Wing of Anopheles
A. maculipennis, Meigen. This species and A. bifurcatus maculipennis, Meigen.
are malaria carriers. True Anopheles only occur in
Europe, North America, the North of Africa and in the mountains of India, and one has
been found by Bancroft similar to A. bifurcatus in Queensland. They are easily told by
the absence of scales on thorax and abdomen, and by the rather densely scaled wings
with lanceolate scales.

Genus. Myzomyia, Blanchard; Grassia, Theobald.

Comp. rend. heb. Soc. Biol., No. 23, p. 795, Blanchard; “Mono. Culicid.,” 1910,
iii, p. 24; v, p. 16, Theobald.
This genus occurs in Asia, Africa and South America, Europe and East Indies. The type
is M. funesta, Giles, found in Central and West Africa. Although structurally there is not
much difference between this genus and Anopheles, they differ greatly in appearance,
and there are usually a few narrow curved thoracic scales projecting over the head,
whilst the wing scales are much smaller in proportion, and the wings more uniformly
spotted, always so along the costa. Funesta and lutzii are undoubtedly malaria bearers
and also rossii.
 Genus. Neomyzomyia, Theobald.
“Mono. Culicid.,” 1910, v, p. 29.
A single species only occurs in this genus, N. elegans, James, from India. In this
genus, which is near to Myzomyia, the fourth long vein is very near the base of the
third, and there are outstanding scales on the prothoracic lobes, and there is a marked
tuft of dense scales at the posterior angles of the head.

Genus. Cycloleppteron, Theobald.

“Mono. Culicid.,” 1903, ii, p. 312; 1903, iii, p. 58; 1910, v, p. 33.
Two common species only occur in this genus, C. grabhamii, Theob., from Jamaica,
and C. mediopunctatus, Theob. (Lutz., ms.), from South America. The chief character is
the presence of large black inflated pyriform scales on the wings. The palpi are densely
scaled. Neither have been shown to be malaria bearers.

Genus. Feltinella, Theobald.

“Mono. Culicid.,” 1907, iv, p. 56.
A single species, so far only found in this genus. The basal lobes of the male genitalia
of two segments, the prothoracic lobes with dense outstanding scales.
The species, F. pallidopalpi, Theob., occurs in Sierra Leone.

Genus. Stethomyia, Theobald.

“Mono. Culicid.,” 1903, iii, p. 13; 1907, iv, p. 59; 1910, v, p. 35.
Four species occur in this marked genus—one S. nimba, Theob., from British Guiana
and Para, another S. fragalis, Theob., from the Malay States, S. culiciformis, James and
Liston, from India, and S. pallida, Ludlow, from India.
The former may be a malaria carrier, for Dr. Low says: “Malarial fever is got amongst
the Indians and often of a severe type. In that connection it is interesting that in the
interior, at a place called Corato, I got an entirely new Anopheles in large numbers.”
The genus is easily told by its unornamented wings, flat head scales, mammillated
prothoracic lobes and long thin legs.

Genus. Pyretophorus, Blanchard; Howardia, Theobald.

Compt. rend. heb. Soc. Biol., No. 23, p. 705, Blanchard; Journ. Trop. Med., v,
p. 181; and “Mono. Culicid.,” 1903, iii, p. 13; 1910, v, p. 36, Theobald.
Forty-four species come in this genus, of which Anopheles costalis, Loew, is the type.
This genus is found in Africa, India, Europe and in Australia. Three species are proved
malaria bearers, namely, P. costalis, Loew, P. chaudoyei, Theob., and P. superpictus,
Grassi. Members of this genus can be told by having narrow curved thoracic scales,
hairy abdomen, and much-spotted wings.

Genus. Myzorhynchella, Theobald.

“Mono. Culicid.,” 1907, iv, p. 78.
In this genus the thorax has distinct, narrow curved scales, and the abdomen is hairy,
the wing scales broad and lanceolate, and the head with broad scales not closely
appressed, but not forked or fimbriated.
Five species are known: lutzi, Cruz; parva, Chagas; nigritarsis, Chagas; tibiomaculata,
Neiva; gilesi, Neiva; and nigra, Theobald. They are all recorded from Brazil, and nigra
also from Mexico.

Genus. Manguinhosia, Cruz, in Peryassu.

“Os Culicideos do Brazil,” 1908, p. 112.
A single marked species from the Brazils. The thorax has piliform curved scales, and
some narrow curved and flattened ones on the sides. Abdomen pilose, except the last
three segments which are scaled. No tufts of scales on posterior femora.
Allied to Lophoscelomyia, but at once told by the absence of scale tufts on the hind
femora. M. lutzi, Cruz, Brazil.

Genus. Chrystya, Theobald.

“Rep. Sleeping Sickness, Roy. Soc. Eng.,” 1903, vii, p. 34.
A very marked genus in which the hairy abdomen has very long, dense, hair-like,
apical, scaly tufts to the segments. A single species only so far known, C. implexa,
Theobald, from Africa (Uganda, Sudan, etc.).

Genus. Lophoscelomyia, Theobald.

Entomologist, 1904, xxxvi, p. 12.
A single species only, from the Federated Malay States. The hind femora have dense,
apical scale tufts; the thorax long, hair-like curved scales; abdomen pilose, except the
last two segments which are scaly; wings with broad, blunt, lanceolate scales.
 Genus. Arribalzagia, Theobald.
“Mono. Culicid.,” 1903, iii, pp. 13 and 81; and 1910, v, p. 48.
Two species only occur, found in South America. The thorax and abdomen have scales
and hairs respectively, as in Pyretophorus, but the abdomen has in addition prominent
lateral apical scale tufts to the segments and a scaly venter. Wings with membrane
tinged in patches and wing scales bluntly lanceolate and very dense. The type is A.
maculipes, Theob. found in Trinidad and Brazil; A. pseudomaculipes, Cruz, also in
Brazil.

Genus. Myzorhynchus, Blanchard; Rossia, Theobald.

Compt. rend. heb. Soc. Biol., 1902, No. 23, p. 795, Blanchard; Journ. Trop.
Med., 1902, p. 181, Theobald; “Mono. Culicid.,” 1903, iii, p. 84; 1907, iv, p. 80;
1910, v, p. 49.
A very marked genus of large, dark, densely scaled species, found in Europe, Asia,
Africa and Australia. The thorax with hair-like curved scales; the abdomen with ventral
and apical scales, and a median ventral apical tuft, and with very densely scaled palpi
in the female, and densely scaled proboscis. It seems to be mainly an Asiatic and East
Indian genus, but three species occur in Africa and one in Australia. They are mostly
sylvan species and bite severely.
Fourteen species are known. Five are malaria carriers (vide list, p. 566).

Genus. Nyssorhynchus, Blanchard; Laverania, Theobald.

“Mono. Culicid.,” 1910, iii, p. 14; v, p. 55, Theobald; Compt. rend. heb. Soc.
Biol., No. 23, p. 795, Blanchard.
A group of small, closely allied species found in Asia, Africa and Australia, twelve out of
the twenty species coming from India.
The thorax is covered with narrow curved and spindle-shaped scales, abdomen with
small, flat or narrow curved dorsal scales, especially on the apical segments or in
patches; the legs are always banded or spotted with white, and the tarsi have as a rule
one or more pure white segments. (This banding and spotting is of no generic value,
however.)
The species show considerable seasonal variation. The type of the genus is N.
maculatus, Theobald.
Three are malaria carriers (vide list, p. 566).
 Genus. Cellia, Theobald.
“Mono. Culicid.,” 1903, iii, p. 107; 1910, v, p. 67.
Very marked Anophelines, with densely scaly abdomens, the scales irregularly disposed
on the dorsum and forming dense lateral tufts; thorax with flat spindle-shaped scales;
palpi densely scaled and also the wings.
The type of the genus is the African C. pharoensis, Theob. It is represented in Asia by
C. kochii, Dönitz; in West Indies and South America by C. argyrotarsis, Desvoidy, and
C. bigotii, Theob.; in Africa by C. squamosa, Theob., etc.
C. argyrotarsis, Desvoidy, and C. albimana, Wiedemann, are undoubtedly malaria
bearers.

Genus. Neocellia, Theobald.

“Mono. Culicid.,” 1907, iv, p. 111.
Allied to Cellia, but has no lateral scale tufts. Three species recorded from India.

Genus. Kertészia, Theobald.

“Ann. Mus. Nat., Hung.,” 1905, iii, p. 66.
This genus has the thoracic scales hair-like, except a few narrow curved ones in front;
abdominal scales long, broad and irregular.
A single species, K. boliviensis, Theob. from Bolivia.

Genus. Manguinhosia, Cruz.

The thorax has narrow hair-like curved scales and some broad straight scales; others
spatulate on the sides. Abdomen with fine hairs, except the last three segments which
are scaly. Tufts of scales on the hind femora. Wing scales lanceolate.
The type is M. lutzi, Cruz, from Brazil.

Genus. Chagasia, Cruz.

“Brazil-Medico,” 1906, xx, pp. 20, 199.
This genus can at once be told by the antennal segments having many dense scaly
tufts. Type, C. fajardoi, Lutz, from Brazil.

Genus. Calvertina, Ludlow.

 Canadian Entomologist, 1909, xli, pp. 22, 234.
The antennæ in this genus have outstanding scales on the second segment, more
appressed ones on the first. At least one abdominal segment with long, flat, more or
less spatulate scales. Type, C. lineata, Ludlow, from Philippine Islands.

Genus. Birónella, Theobald.

“Ann. Mus. Nat. Hung.,” 1905, iii, p. 69.
At once told by the first submarginal cell being very small. Type, B. gracilis, Theob.
from New Guinea.

Sub-family. Megarhininæ.
Three genera occur in this marked sub-family; they are the largest of all mosquitoes,
and are very brilliantly coloured, and many have tail fans. They occur in North and
South America, Asia, Africa, and Australia. The long curved proboscis is very marked.
They are usually spoken of as elephant mosquitoes; some are vicious blood-suckers at
times.
The three genera tabulate as follows:—
α. Palpi long in both sexes.
β. Last segment of ♀ palp round or Genus Megarhinus, R.
blunt as if broken Desvoidy.
ββ. Last segment of ♀ palp long Genus Ankylorhynchus,
and pointed Lutz.
αα. Palpi of female short of male
long.
Palpi of female not more
than one-third length of Genus Toxorhynchites,
proboscis Theobald.

Genus. Megarhinus, Robineau Desvoidy.

“Mém. Soc. d’Hist. nat. de Paris,” 1827, iii, p. 412; “Mono. Culicid.,” 1901, i,
p. 215; 1903, iv, p. 163; 1907, iv, p. 128; 1910, v, p. 89.
All large brilliant mosquitoes with long palpi in both sexes and, as a rule, with a caudal
fan of scales; the proboscis is long and bent. They are all sylvan species, and are not
so far recorded as biting man.

Genus. Toxorhynchites, Theobald.

 “Mono. Culicid.,” 1901, i, p. 244; 1903, iii, p. 119; 1907, iv, p. 140; 1910, v, p. 95.
Differs from the former genus in that the female palpi are short. The palpi may have
one, two or three minute terminal segments. Banks’s genus Worcesteria has three.
The elephant mosquito of India (T. immisericors), Walker, bites very severely. They are
sylvan species.

Sub-family. Culicinæ.
Genus. Mucidus, Theobald.
“Mono. Culicid.,” 1901, i, p. 268; 1910, v, p. 125.
This genus is so far confined to Australia, West and Central Africa, India, East Indies
and Malay Peninsula. They are all large mosquitoes, easily told by the whole body
being more or less covered with long twisted scales, giving them a mouldy appearance,
and the legs densely scaled with outstanding scales; the wings with large parti-
coloured scales. The Australian M. alternans, Walker, occurs in larval form both in fresh
and salt water. The adults bite man.

Genus. Psorophora, Robineau Desvoidy.

“Mém. de la Soc. d’Hist. nat. de Paris,” 1827, iii, p. 412, R. Desvoidy; “Mono.
Culicid.,” 1901, i, p. 259; 1903, iii, p. 130; 1907, iv, p. 158; 1910, v, p. 123,
Theobald.
This genus is confined to the Americas and the West Indies. Several species exist
which can easily be told from Mucidus by the absence of long twisted scales and the
narrower wing scales. The legs are densely scaled and the thorax ornamented with flat
spindle-shaped scales.
P. ciliata, Robineau Desvoidy, occurs in both North and South America, and bites man.

Genus. Janthinosoma, Arribalzaga.

“Dipt. Arg.,” 1891, p. 52, Arribalzaga; “Mono. Culicid.,” 1901, i, p. 253; 1903, iii,
p. 124; 1907, iv, p. 152; and 1910, v, p. 118, Theobald.
Hind legs only densely scaled; some of the hind tarsi are always white. The venation is
as in Culex. The abdomen is metallic and iridescent. They all bite man and occur only
in the Americas and West Indies.

Genus. Stegomyia, Theobald.

 “Mono. Culicid.,” 1901, i, p. 283; 1903, iii, p. 130; 1907, iv, p. 170; 1910, v,
p. 151.
This, the most important genus in the Culicinæ, can be told by the head and scutellum
being clothed with flat scales and the thorax with narrow curved ones.
About forty species are known in this genus, occurring in Southern Europe, Asia, Africa,
Australia, the Americas, East and West Indies, and on most oceanic islands. Many of
them seem to be vicious blood-suckers. They are mostly black and white mosquitoes,
and several seem to go by the name of tiger mosquitoes. The genus contains the
yellow fever mosquito (S. fasciata, Fabricius), the only one that need be dealt with in
detail here. The chief known species tabulate as follows:—
 A. Proboscis banded.
α. Legs basally banded.
Thorax brown, with scattered creamy-white
scales annulirostris, Theobald.
Thorax black, with narrow, curved golden
scales periskelta, Giles.
αα. Legs with basal and apical banding. Fore legs
with no bands; mid with apical and basal
bands on first and second tarsals, hind
with basal bands.
Thorax white in front, with a brown eye-like
spot on each side thomsoni, Theobald.
AA. Proboscis unbanded.
β. Legs basally banded.
γ. Abdomen basally banded.
Thorax with one median silvery-white line scutellaris, Walker.
pseudoscutellaris,
Thorax as above, but pleuræ with white lines
Theobald.
Thorax similar, but two white spots near
where line ends gelebinensis, Theobald.
Thorax with two median yellow lines and
lateral curved silvery lines fasciata, Fabr.
Thorax with two short median lines and a
white patch on each side nigeria, Theobald.
Thorax with large lateral white spots in front,
smaller ones by wings, two narrow median
lines and two posterior sub-median white
lines lilii, Theobald.
Thorax with a white W-shaped area in front,
a prolongation curved on each side
enclosing a brown eye-like spot W-alba, Theobald.
Thorax with white frontal median spot, two
large lateral spots, a small one in front of
the wings, a narrow median white line and
narrow sub-median ones on posterior half.
Last two hind tarsi white wellmannii, Theobald.
Thorax brown, with broad white line in front
extending laterally towards wings, where
they swell into a large patch, a white line
on each side just past wing roots. Last two
hind tarsi white albipes, Theobald.
Thorax with silvery white spot on each side
in front, small one over roots of wings and
white over their base Last two hind tarsi
 white over their base. Last two hind tarsi
white pseudonigeria, Theobald.
Thorax with two lateral white spots, front
ones the largest, a small median one near
head, two yellow median lines, a short
silvery one on each side before the
scutellum simpsoni, Theobald.
Thorax with silvery-white scaled area in front argenteomaculata,
and another on each side in front of wings Theobald.
Thorax with median yellowish-white line, a
silvery patch on each side in front of wings
extending as a fine yellow line to the
scutellum, and another silvery spot before
base of each wing poweri, Theobald.
Thorax with small grey-scaled area in front
of wing roots and three short creamy lines
behind minutissima, Theobald.
Thorax (?) denuded; abdomen black; fifth
segment with yellow basal band; sixth
unbanded; seventh, two median lateral
white spots; eighth, two basal lateral white
spots; second hind tarsal nearly all white dubia, Theobald.
γγ. Abdomen unbanded.
First hind tarsal all white, second basally
white, last two dark. Thorax chestnut
brown, with a broad patch of white scales
on each side in front and a median pale
line terreus, Walker.
ββ. Legs with white lines as well as basal bands.
Thorax brown, with white lines; abdomen
with basal bands grantii, Theobald.
Fore and mid legs with apical bands, hind
βββ.
basal.
Fourth tarsal of hind legs nearly all white mediopunctata, Theobald.
Mid metatarsi with basal pale banding, base
and apex of hind, also base of first tarsal
pale assamensis, Theobald.
ββββ. Legs unbanded.
δ. Abdomen basally banded.
Thorax with front half white, rest bronzy-
brown pseudonivea, Ludlow.
Thorax deep brown, with scattered golden
scales, showing two dark eye-like spots;
head white, dark on each side and behind albocephala, Theobald.
Thorax brown with golden stripes; abdomen
 Thorax brown with golden stripes; abdomen
with narrow basal bands on fifth and sixth
segments only auriostriata, Banks.
δδ. Abdominal banding indistinct.
Thorax with broad silvery white patch on
each side in front albolateralis, Theobald.
δδδ. Abdomen unbanded.
argenteopunctata,
Thorax with six silvery spots
Theobald.
δδδδ. Abdomen with apical white lateral spots.
Thorax unadorned, except for pale scaled
lines laterally punctolateralis, Theobald.
δδδδδ. Abdomen with basal white lateral spots.
Thorax with two pale median parallel lines
and two silvery lateral spots ininuta, Theobald.
Thorax unadorned.
A white spot middle of head tripunctata, Theobald.
No white spot amesii, Ludlow.
AAA. Proboscis yellow basally, dark apically.
Abdomen with apical pale bands crassipes, Van der Wulp.
Proboscis with median interrupted white line on
AAAA.
basal half.
Head black, anterior margin grey albomarginata, Newstead.
Stegomyia fasciata, Fabricius (Yellow Fever Mosquito).
This insect, which is the proven carrier of yellow fever, is commonly called the tiger,
brindled, spotted day or striped mosquito. It is also referred to by some writers as S.
calopus, Meigen. It is subject to considerable variation in colour, but the thoracic
markings are generally very constant. The general colour is almost black to deep
brown, the head with a median white area, white at the sides and in front around the
eyes; the thorax has two median parallel yellow lines, a broad curved silvery one on
each side and white spots at the sides; the scales on the intervening spaces of the
thorax are brown. The dark abdomen has basal white bands and basal white lateral
spots. The dark legs have basal white bands, the last segment of the hind legs being
all white except in a variety from South America and the West Indies (luciensis), which
has the tip of the last hind tarsal dark. The abdomen may also vary in colour, some
having pale scales over most of the surface (queenslandensis).
The food of the adult female consists mainly of man’s blood, but she will also feed on
dogs and other animals. The male has been said to bite, but such is very unusual. This
mosquito bites mainly in the daytime up till about 5 p.m.
The adults breed the first day after emergence. They may live a considerable time,
Bancroft having kept females for two months in confinement. The ova are laid
separately, often in chains; they are black, oval, with a reticulated membrane outside,
some of the reticulated cells containing air. They may hatch in from six to twenty
hours, the larval stage nine days, the pupal stage three; thus the whole cycle may be
completed in from twelve to thirteen days. The ova when dry can remain undeveloped
for a considerable time. The larvæ are greyish-white, with short, thick siphon, and feed
at the bottom of the water, only coming to the surface now and again to breathe. This
is almost entirely a domesticated gnat, seldom being found far from man’s habitations.
Its larvæ occur in such small collections of water as old sardine tins, jam-pots,
calabashes, puddles, barrels, wells—in fact, wherever water is held up, even to the
gutters of houses. Not only are they found breeding on land, but also on board ship,
although they prefer artificial collections of water. They may also breed in larger
natural collections.
This insect is easily transported by steam and sailing ships and by train, and this
doubtless explains its very wide distribution. The adults may live for fifty days, and it is
on this account and their frequent occurrence on ships that danger lies in regard to the
Panama Canal. An infected insect may leave that endemic centre of yellow fever and
live until the vessel arrived at the Philippine Islands and fly ashore, and so introduce
the disease for the native fasciata possibly to spread.
Roughly the distribution of this pest is as follows: Africa from South to North, but
especially along the coast and up the Nile. In Asia, in India, Ceylon, Burma, Siam,
along the ports of the Malay Peninsula, in French Cochin China, Philippine Islands, the
Andaman and Nicobar Islands, Japan, Malay Archipelago, and East Indies, Turkey in
Asia, Arabia and Palestine.
In Australia it occurs in Queensland, New South Wales, Victoria and South Australia.
In Europe in Italy, Spain, Portugal, Greece, in the Mediterranean Islands.
In South America, Central America, Mexico, North America, and the West Indies it is
very abundant, and it also is found in the Bahama Islands, Fiji, Sandwich Islands,
Samoa, the Azores, Teneriffe and Santa Cruz, Pitcairn Islands and Bermuda.
For a full account of its distribution the reader is referred to the following: “The
Distribution of the Yellow Fever Mosquito (Stegomyia fasciata, Fabricius) and General
Notes on its Bionomics;” “Mém. 1er Congrès international d’Entomologie, 1911, ii,
pp. 145–170, F. V. Theobald.” In addition to being the yellow fever carrier, it is
supposed by Wenyon to be the intermediate host of the parasite of Bagdad sore.

Stegomyia scutellaris, Walker.

A vicious biter, found in India, China, Malay, East Indies, and Ceylon. The thorax has
one median silvery stripe, and so can easily be told from S. fasciata.
A very similar species occurs in Fiji, but can be told by the pleuræ having white lines,
not spots (S. pseudoscutellaris, Theobald). It is the intermediate host of filaria in Fiji
(Bahr).
A number of nearly allied genera occur here (vide synoptic table).

Genus. Theobaldia, Neveu-Lemaire.

Theobaldinella, Blanchard.
Includes several large Culicines, of which T. annulata, Meigen, is the type. The wings
are usually spotted (annulata, incidens, etc.), but may be nearly plain (spathipalpis).
The males have the palpi swollen apically, and the females have long five-jointed palps.
Several of these are vicious biters.

Theobaldia annulata, Meigen.

This large gnat (6 mm. long) can be told by its wings having five large spots of dark
scales and by its legs having broad basal white bands to the tarsi. The larvæ occur in
rain barrels and small pools. It is essentially a domestic form, occurring in houses and
privies. Its distribution is Europe generally and North America. The bite is very severe,
and in some districts gives rise to painful œdema. 392
Theobaldia spathipalpis, Rondani, occurs in Italy, Mediterranean Islands, Palestine, the
Himalayas, Khartoum, and in South Africa. It is about the same size as T. annulata, but
is yellowish-brown in colour, with striped thorax and mottled and banded legs. It
occurs in privies and bites very severely.

Genus. Culex, Linnæus.

“Syst. Nat. Ed.,” 1758, x, Linnæus; “Mono. Culicid.,” 1901, i, p. 326; 1910, v, p. 322,
Theobald.
This large genus still contains many forms which
should be excluded. The species normally have
narrow curved median head-scales, and similar ones
on the scutellum; the female palpi are shorter than
Fig. 400.—Wing of a Culex.
in the former genus and the male palpi are pointed;
the lateral vein-scales are narrow and linear.
The type is Culex pipiens, Linn., the common gnat of Europe. The thorax is covered
with narrow curved golden-brown scales, the abdomen has basal pale bands to the
segments and the legs and proboscis are unbanded. The stem of the first submarginal
cell is always less than one-fifth the length of the cell. It lays its eggs in rafts in water-
butts, etc., and even in the foulest water. They are first deposited in England in June
and July, and again soon after hatching in August. In some districts this gnat bites man
viciously, in others not at all.
The common tropical gnat (Culex fatigans, Wied). This resembles the European Culex
pipiens, but can always be told by the stem of the first submarginal cell always being
much longer than it is in C. pipiens. This is one of the species that has been proved to
transmit filariæ to man, etc. Varieties of it occur in almost every country between 40°
N. and S., having a very similar range to S. fasciata. In all countries it appears to be
connected with the transmission of Filaria bancrofti, and it is also said to carry the
micrococcus of dengue fever.

Genus. Melanoconion, Theobald.

“Mono. Culicid.,” 1903, iii, p. 238; 1907, iv, p. 507; 1910, v, p. 455.
This genus is composed of eight species, most of which are small black gnats which
bite viciously and which occur in swamps and jungles. They can at once be told from
Culex by the veins of the wings having dense broadened scales on their apical areas
and along the upper costal border. The femora and apices of the tibiæ are swollen.
The black mosquito, Melanoconion atratus, Theob. This small gnat is a very
troublesome pest in swamps in the West Indies. The female bites both by day and by
night, and the bite causes severe irritation. The larvæ live in permanent ponds. It is
almost black in colour, but sometimes presents a dull coppery sheen; each segment
has small lateral basal white spots. Length 2·5 to 3 mm.
It occurs in Para and British Guiana as well as in the West Indies.
Ordinary mosquito netting is no use for keeping off this pest.

Genus. Grabhamia, Theobald.

“Mono. Culicid.,” 1903, iii, p. 243; 1907, iv, p. 284; and 1910, v, p. 277.
Allied to Culex, but separated by the wings having short fork-cells, mottled scales, the
median ones thick and also some of the lateral ones short and broad; the last two
joints of the male palps are very slightly swollen. The eggs are laid singly, not in rafts,
and the larvæ have short, thick siphons. Ten species occur and are found in Europe,
North America, West Indies and Natal. G. dorsalis, Meigen, bites severely in Europe. G.
sollicitans, Walker, is a great scourge along the New Jersey Coast and at Virginia
summer resorts and in Florida. It breeds in brackish water and is the most common
mosquito of the Atlantic seaboard.
 Genus. Pseudotæniorhynchus, Theobald;
Tæniorhynchus, Theobald, non-Arribalzaga.
Differs from the former in having the whole wing veins clothed with dense, broadish
elongated scales. They occur in South America (T. fasciolatus, Arri.), in Africa (T. tenax,
Theob.), in Europe (T. richardii, Ficalbi). The latter bites very severely.

Genus. Tæniorhynchus, Arribalzaga; Mansonia, Blanchard;

Panoplites, Theobald.
Compt. rend. heb. Soc. Biol., 1901, iii, 37, p. 1046; “Mono. Culicid.,” 1901, ii,
p. 173; and 1910, v, p. 446, Theobald.
A very marked genus, easily told by the broad asymmetrical wing scales. It occurs in
Africa (T. africana and T. major, Theob.); in Asia (T. uniformis, Theob.; T. annulipes,
Walker, etc.) and in Australia (T. australiensis); in the Americas and West Indies (T.
titillans, Walker). The eggs (fig. 395, d) are peculiar in form and are laid separately;
the larva has not been described; the pupa has long curved siphons. They mostly occur
along rivers, in swamps and forests, and bite very severely. They also enter houses (T.
titillans). T. uniformis is most troublesome during the rains. The saliva is strongly acid.
Both these species carry the larvæ of Filaria bancrofti.

Genus. Chrysoconops, Goeldi.

“Os Mosq. no Para,” 1905, p. 114, Goeldi; “Mono. Culicid.,” 1910, v, p. 433, Theobald.
Bright yellow or yellow and purple mosquitoes, with rather dense wing scales.
Numerous species occur in Africa (aurites, annettii, fuscopennatus, etc.), others in
India, Australia and South America.
Low found filariæ in the thoracic muscles of fuscopennatus in Uganda.
Several of the Ædeomyina bite, especially the small Uranotænias. They are all sylvan
species, seldom entering houses. They need not, therefore, be referred to here.
For full details of the Culicid genera and species the reader is referred to my
monograph 393 and other works mentioned below.

Other Nematocera.
Other nematocerous flies are midges, daddy-long-legs and sand-flies. The ones which
cause annoyance to man besides Culicidæ are the following:—
Sand-flies (Simulidæ), certain midges (Chironomidæ), and a few owl midges
(Psychodidæ).
The Nematocera have long thread-like jointed antennæ and their pupæ are, as a rule,
naked; the larvæ have a distinct head and can thus be told from the next section
(Brachycera).

Family. Simulidæ.
This family consists of a single genus, Simulium, Latreille, which Roubaud has
recently divided into two sub-genera called Pro-Simulium and Eu-Simulium. These
insects, which are frequently spoken of as sand-flies, are found in all parts of the
world; they are all small insects varying from 1·5 to 3 mm. The females are very
bloodthirsty, but the males appear to be incapable of sucking blood.
The head sunk under the humped thorax; antennæ short, straight; palpi short and
broad, of four segments, bent; wings broad and in some iridescent, legs stout. The
male has holoptic eyes, whilst in the female they are small and widely separate. The
sucking proboscis is short. The thorax and abdomen are clothed with short hairs
which may form spots and markings; these are golden, silvery, grey, or brownish. In
the sub-genus Pro-Simulium the second segment of the hind tarsi in both sexes is
elongate, linear, and without a basal notch; in Eu-Simulium it is short, curved, and
dorsally notched at the base.
Simulidæ often occur in swarms, and attack not only man but cattle, horses, and
poultry. In some districts they are more annoying than mosquitoes.
Their life-cycle has been most completely worked out by King, in Africa.
The larvæ and pupæ occur in swiftly flowing water, by waterfalls, in rapids, etc. The
ova are laid in gelatinous masses on plants or rocks close to or overhanging the
water. The larva is cylindrical, enlarged posteriorly, where it is provided with a sucker,
by means of which it attaches itself to a rock, water weeds, debris, etc.; anteriorly it
has a proleg close behind the head on the lower surface. The head is dark and
chitinous. The respiration takes place by means of branched tracheal gills which
protrude from the dorsal surface of the last body segment; they are retractile. The
colour varies from deep green to yellow or almost black. Their food consists of algæ
and other organisms in the water brought to their mouth by two fan-like organs
placed on the head. The larvæ can crawl from place to place by means of the thoracic
proleg; they occur in masses, usually in a more or less erect attitude. A network of
threads is spun on their support, by means of which King tells us “they are enabled to
maintain their position against the strongest current; frequently they will leave their
support and let themselves out into the stream anchored by threads of silk and
enabled by them to return.”
When full fed the larva spins a pocket-shaped cocoon on the support, within which it
pupates. The pupa is motionless and has a pair of branched spiracles projecting from
behind the head. When the adult emerges, a bubble of air collects around it, and in
this it floats to the surface and at once takes wing. The European species take a
month to complete larval life, a week being spent in the pupal stage. The flies are
most restless, and even when stationary continually move their legs about like
feelers. Sometimes the swarms consist entirely of females, sometimes early in the
season mostly of males.
The females pierce the skin of humans on tender spots, such as ears, the forehead,
around the eyes and nose, and crawl into the cavities. They are quite harmless at
night, mainly attacking about sunrise and sunset. Some crawl up the arms and legs
and down the neck, and leave behind little red weals which itch intensely (S.
damnosum, Theob.), and blood may flow freely from the wounds.
The following are some of the worst species:—
Simulium columbaschensis, the “Kolumbatz fly,” which abounds in the damp marshy
lands along the Danube, and is a great plague to man and beasts in Hungary, and is
also abundant in Austria and Moravia, and is most numerous after inundations from
the Danube. They sometimes appear in such swarms that it is impossible to breathe
without getting them into one’s mouth. There are instances of children being killed by
these flies when left on the ground by their mothers when working in the fields.
S. damnosum, Theob. This occurs throughout Equatorial Africa and is known as the
“jinja fly” in Uganda, the “fouron” in the French Congo, the “kilteb” in the Sudan. It is
a most vicious biter, and in some parts occurs in “belts”; Dr. Christy found one such
extending from the shores of the Victoria Nyanza northwards along the right bank of
the Nile for twelve or fifteen miles or more, and perhaps three or four miles wide. In
this area the flies swarm in millions at certain seasons, so much so that the natives
have to leave their plantations. The bite causes a weal, marked by a drop of blood.
S. griseicollis, Becker. The so-called “nimitti” occurs in Upper Egypt and the Anglo-
Egyptian Sudan. It lives near the river and is not found more than half a mile from it.
Human beings are bitten on the face and hands, animals in the region of the
pudenda.
S. latipes, Meigen. This is a European species, also found in Natal.
S. wellmanni, Roubaud. The “ohomono” of Angola, where it bites viciously and is
dreaded by the naked porters.
S. buissoni, Roubaud. Occurs in abundance in the Marquesas Islands. It has been
suggested that this species may help to propagate leprosy. 394
A large number of these insects have been described by Lutz in Brazil. 395
A Simulium sp. (?) is very harmful to poultry in Cape Colony. 396
In America, Simulidæ are most annoying. One, S. meridionale, Riley, also known as
the turkey gnat in the Mississippi Valley, has been supposed to be the carrier of
chicken cholera; anyhow, it has caused the death of thousands of chickens and
turkeys in Virginia annually. 397
In Mexico Townsend found a Simulium which was named S. occidentalis, which
caused great annoyance to man, many people being so susceptible to them as to
preserve through the gnat season a chronic inflammation of the exposed parts of face
and neck, resulting from the repeated bites giving rise to sores. 398
Men and horses have been partially incapacitated by the bites of sand-flies or
Simulium in a Hampshire wood (Cantlie, Brit. Med. Journ., April 28, 1900, v, No.
2,052, p. 1023).

Fig. 401.—Wing of Fig. 402.—Wing of

Simulium. Chironomus.

Family. Chironomidæ (Midges).

The Chironomidæ or midges are not only frequently mistaken for mosquitoes, but
some are very annoying to man by biting him as mosquitoes do. They are easily
distinguished from true mosquitoes (Culicidæ) by the following characters: (1) head
small, often retracted under the cowl-like thorax; (2) no scales to the wings or body;
and (3) the different arrangement of veins on the wings (fig. 402).
Two genera are important as annoying man, namely, Culicoides, Latreille, and
Johannseniella, Williston. The larvæ of Chironomidæ are either aquatic, both fresh
water and marine, and help to make the former foul, 399 according to Slater, or may,
as in Ceratopogoninæ, live beneath the bark of trees, etc. The pupæ are very varied
and also the life-histories of the different genera. 400 The blood-sucking habit is
confined to the sub-family Ceratopogoninæ.

Sub-family. Ceratopogoninæ.
This sub-family of midges consists of very small species varying from 1 to 2 mm. in
length; the wings have darkened areas, and the second longitudinal vein is wanting,
and the first and third veins are stouter than the others and placed close to the
anterior margin, the fourth and fifth are forked; the antennæ in both male and female
are composed of fourteen segments, six or eight in the males bearing long hairs.
The chief blood-sucking species belong to the genera Culicoides, Latreille, and
Johannseniella, Williston. The latter genus differs from the former in the absence of
an empodium or median appendage on the last segment of the tarsi. The genus
Ceratopogon, as restricted by Kieffer, is not supposed to take vertebrate blood, but
Austen has recently noticed that the type specimen of C. castaneus, Walker, and a
new species described by him, apparently have their bodies distended with blood.
The wings in the Ceratopogoninæ are carried flat when at rest.

Fig. 403.—A Ceratopogon, or midge. Greatly

enlarged.

In spite of their small size the females are the most bloodthirsty and annoying of all
insects. The Culicoides, which are often called “sand-flies,” bite during the day and
rarely at night. Usually they are most troublesome between 3 and 6 p.m. They
frequently attack in swarms, especially in the open, and owing to their minute size
can get through fine mosquito netting. Some of them produce a distinct “buzz” when
on the wing. These insects are found in all parts of the world. No species has been
definitely connected with any disease, but Culicoides has been suspected of carrying
the germs of Delhi boil. The larvæ of Culicoides are elongate in form and have
smooth bodies composed of thirteen segments including the head, which is horny;
there is no proleg on the first segment as seen in Chironomus, and on the anal
segment are retractile gills. They are very active and live in the sap of various trees
which saturates diseased bark.
The pupæ are smooth, but the abdominal segments bear a transverse row of small
spines. Austen describes a number of Culicoides and one Johannseniella and three
Ceratopogons from Africa, 401 and Lutz 402 a number of this sub-family from Brazil,
including a new genus, Centrorhyncus. Another genus, Tersesthes, Townsend
(Centrotypus, Grassi; Mycterotypus, Noe), also occurs in Brazil.
Culicoides ornatus, Taylor, is described from Townsville, Australia, found in mangrove
swamps. It is a very vicious biter and causes considerable irritation, settling on hands
and wrists (Taylor, Rep. Ent. Aust. Inst. Trop. Med. (1912), 1913, p. 24).
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