ON THE IDENTITY OF CULEX PIPIENS LINN/EUS.
[Diptera, Culicidae.]
[PLATESI TOIII.]
BY
HARRISON G.DYAR
and FREDERICKKNAB.The
family Culicidse, comprising the mosquitoes, is basedupon
thegenusCitle.r of Linnaeus,ofwhich
the generally recog- nized type isCulex
pipiens Linnaeus.The
genusand
species datefrom
the beginning of binomial nomenclature, the tenth edition of theSystema
Naturae. In this work, page 602, Lin- naeus treats thegenus
thus:VI.
DIPTERA.
224 CULEX. Os
aculeis setaceis intravaginam
flexilem.pipiens. 1. C. cinereus, abdomine annulis fuscis octo. Fn. svec. 1116.
Fl. Lapp. 363, 364.
Blank ins. t. 15. /. A-D.
Reaum. ins. 4. t. 43, 44.
Swamm.
quart, t. 23.bibl. t. 31. /. 4-8. /. 32. /. 1-5.
Joblot. micr. 1. pp. 2. t. 13. /. A. B. C. D. E. H. I. L.
ft. Kalm. itin. 2. p. 268. Musquetoes.
Habitat in Europae aquosis; copiosissima in Lapponia;
ctiam in America obvia. Kalm.
Mas
antennis plumosisvix pungit out sugitsanguinem.Larva in Aquis; Pupa bicornis reversal Insectum pipiens, pungcns; in Indiis magis venenata. An- scres allicit, pullos GaUinarum nutrit
Lapponum
calamitas felicissima.
bifurcatus. 2. C. fuscus, rostro bifurco. Fn. svec. 1115.
Raj.ins. 74. .4.
Reaum. ins. 4. t. 40. /. 1, 2.
Habitat in Europa.
pulicaris. 3. C. alishyalinis:tnaculis tribus obscuris. Faun svec. 1117.
Fl. Lapp. 365.
Derh.phys.-th. 1. i. c.u,f, 5,6.
Habitatin Europa; in America. Kalm.
Cursitat, mordet, relinquit punctum fuscum. Amoen.
acad. 3. p. 343.
reptans. 4. C. niger, alis hyalinis, pedibus nigris annulo albo. Fn.
svec. 1118.
Habitat in Europa,reptatu molestus tempore vespertino.
(.quinus. 5. C. ater, abdomine fusco, fronte alba. Fn. svec. 1120.
Fl. Lapp. 359.
Habitat circum Equos, quorum sanguinem haurit, inter crines corum cursitans.
stercoreus. 6. C. testaceus, alis reticulatis, linea thoracis, tribusque abdominis nigricantibus.
Habitat in stercoribus.
OF WASHINGTON,
VOLUMIi XI, 1909. 31As
above stated, Cule.v pipiens has generally been recognized as the type of the genus,and
thatno
doubt correctly, as thiswould seem
to be themost common, and
in a sense, officinal species included,and would
thus be intended as the typeon
Linnaeus'own
ideas.Moreover,
it is the first species.Of
the others, bifnrcatushas been recognized as a mosquito, anEurop-
ean species of Anopheles: but very little is said about it,and
itwas
obviously subordinated in Linnaeus' conception of the group.The
other species are not mosquitoes at all: pulicaris is a Ceratopogon, ref>tansand
eqninns are species of Sitinilinin,and
stercorens is apparentlysome
Cyclorhaphid fly.They do
not agree with Linnaeus' definition
"Os
aculeis setaceis intravaginam
flexilem,''and
he could not havehad any
ofthem
inmind
as typical of the genus.The
only species so agreeingare the first two, of
which
the first ismuch
thecommoner and
better
known. We
therefore seeno
reason for attempting to reverse the generally accepted procedure,and would
confirm Cule.v pipiens Linnaeus as the type of the genusand
family.The
next point to be determined is the identity of Cule.v pipiens. Linnaeus obviously intended the term to cover allmosquitoes
known
to him, directly or indirectly, except the Anopheles, with long palpi in the female,which
he separatesunder
thename
Cnlc.v bifnrcatus. Linnaeushad
experienced mosquitoes in Lapland,and
they evidentlymade
a lasting im- pressionupon
his mind, as he speaks ofthem
as"most abund-
ant"and
as a terrible plague in that country.He
also includes thecommon
domestic forms, as he speaks of their furnishing food to chickens,and
he also refers to their occurrence inAmerica and "The
Indies," but not by personal observation.The
first separation of speciesfrom
this aggregatewas
by Linnaeus himself, in the followingmanner
: Previous to the date of the tenth edition of theSystema
Naturae, Linnaeushad
published Cule.v vulgaris (Act. "Ups., 31, 1736)and
Cule.v alpinus (Flor. Lapp., 364, 1737) ; but as these antedate the beginning of zoological nomenclature, they are excludedfrom
consideration.Moreover,
Linnaeus himself refers these species assynonyms
of pipiens in the second edition of theFauna
Suecica, 1761.
However,
the descriptions reappear in the second edition of the Flora Lapponica, 1702, so thenames may
be considered to
become
valid on that date.Under
ordinaryconditions, the fact that the
names
first appeared assynonyms
would
render their subsequent use inadmissible; but in Lin- naeus' casewe
hold that an exception should bemade, and
the resurrection of thenames
allowed. Linnaeus first proposed thenames
to represent valid species, and subsequently referredthem
to thesynonymy. He had
clearly the right to again resurrect them.The
only reasonwhy
hehad
apparently not that right is because the arbitrarily established beginning of zoological nomenclature, 1758,happened
to intervene be-tween
the proposal of thenames and
their reference to thesynonymy, which
caused their first valid appearance as syn-onyms
withoutaccompanying
description.We
think that in this case the fact of their previous descriptionmay
justly be considered.Of
these species, Cule.v vulgaris is apparently aSimulium,
but Cule.v alpinus is a mosquito,and
judgingfrom
its place of occurrence, anAedes
in the broader sense. This constituted a restriction ofthe original conception of Cule.v,by
theelimina- tion of theAedes
element, theforms abundant
in Lapland,and
leaves for ourconsideration only thecommon
domestic species.We
exclude also the exoticforms
as not personallyknown
to Linnaeus. In this coursewe
followand
confirm theconclusions of F. V.Theobald
in hismonograph
of the mosquitoes of the world,who
recognizes as Cule.v pipiens thecommon
domestic mosquito of Europe.Having
arrived at this point,we
next inquirewhat
is thecommon
domestic mosquito ofEurope and
does it occur inAmerica? We
find inAmerica no
less than five closely allied but distinct species living as larvae in artificial accumulations of waterand
infesting dwellings as adults,which have
beenknown
tous collectively as "Cule.vpipiens, thehouse mosquito."It is probable that there are likewise in
Europe
several speciesmixed under
thename
Cule.v pipiens. Ficalbi (Bull. Soc. Ent.Ital.,
xxvin,
289, 1896) takes as Cule.v pipiens thecommon
house mosquito of Italy and describes it fully.
He
had,how-
ever,
some
doubt as to the correctness of his identificationand
suggests the alternativename
Cule.v ha-inatophagus.Now
his description indicates that the house mosquito of Italy is not the
same
species as the one of northern Europe. This latter should clearly be recognized as the trueCulex
pipiens, while the othermay
bear thename
Cule.v haeinatophagus Ficalbi.Itappearsto be especially distinguished
by
thelateralexpansion of theabdominal bands
on the lasttwo
segments,which
does not occur in the true Cule.vpipiens, norin Cule.v fatigans,which Theobald
recognizesfrom
the Mediterranean region. It sug- gests in this character the American. Cnle.v siinilis Theobald, an Antillean species resembling the pipiensgroup.Theobald
recognizes that Cnle.v pipiens extends toNorth
America,and
IK- separatesunder
thename
CnJe.v fatigans themore
southerly distributed form, extending through the tropicsOF WASHINGTON, VOLUME
XI, 1909.of both*the
new and
old worlds.He
doesnot, however,go
into the matter in sufficient detailand
his presentation is not con- vincing.We
find in themale
genitalia excellent characters of specific definition. It is true thatTheobald
describes these structures, but he has not illustrated with accuracy the basal parts thatshow
the specific distinctions to the best advantage, having confined his attention chiefly to theappendages
of theside pieces,
which
are practically alike in all the species.The
exact definition ofCulex
pipiens, therefore, still remainsin
some
obscurity,and we
proposenow
toresolveit.We would
restrict
Culex
pipiens to that species with the basal clasps of themale
genitalia asshown
in our figure 4, plate n.We
omit the side pieces in our figures as they are alike in all the species referred to.Our
specimens arefrom Copenhagen, Denmark,
(Dr. F. Meinert), St-Remy-la-Varenne,Maine
et Loire, France (R.du
Buysson),and
Buda-Pesth,Hungary
(Dr. C.Kertesz).
We
have bred specimens with similar genitalia in America,and
are able to confirm the occurrence of the species here,no
doubt as an importation.Our common
rain-barreland
house mosquitoes comprise the following, as defined by the male genitalia.The
adults are generally separableon minor
characters of coloration,which we
shall describe fully elsewhere; but the colorational char- acters of these species are unstableand
unreliable to a large extent; neither are the venational differences pointed outby Theobald
ofany
value.Culex pipiens Linnaeus.
The
harpes (Plate n, fig. 4, a) with acrown
of spines asin all the species here discussed, but with rudimentary basal process (fig. 4, b}.
Harpagones
divided into four plates, the upper one (fig. 4, r) rather narrow, long, exceeding the tip of the harpes, often bent nearly at right angles near the middle;second piece (fig. 4, a?) a rather broad plate
narrowed
at the tip, with arounded
point: third piece a thin, often obscure plate, similar to the second piece but smallerand weaker;
fourth piece (fig. 4, /) abroad, concave stout plate with a nar-
row rounded
tip.The
species is especially characterizedby
having the harpa- gones divided into four plates, all the allied species having atmost
three. In the lowestforms
hereshown
the harpones are divided intotwo
separate plates, the outer one variously toothed, as in pro.vinius (fig. 6), salinarius (fig. 7), similisfig. 8s), and lachrimans (fig. 9) : next they
become
divided into three plates, as in restnans (fig. 5), the second division stilltoothed, although in a simpler
manner
; next there are three divisions, as in quinqucfasciatus (fig. 1), 'coniitatus (fig. 2),and
dipscticits (fig. 3), but without teeth,and
finally four in pipicns.Moreover
the plates are ofmoderate
length,none
especially elongated or shortened; all have
rounded
pointedtips, without
any
teeth.We
are not sufficiently acquainted with theEuropean
faunato say
whether
Culc.v pipicns develops local races or allied species in outlying countries adjoining its range, as quinquc- fasciatnsand
siinilis, hereinafter discussed, do; but such isprobably the case. In
America
it does not,and
this furnishes additional support for the opinion that the species has been introduced comparatively recently into America.Culex quinquefasciatus Say.
This
seems
tobe the earliestname
for the widely distributed species called Citle.v fatigansby
Theobald.We have
preparedmounts
of the genitalia of specimensfrom
various parts ofNorth and South
America, Hawaii, the Philippines,and
India,and
findthem
constant. Theobald's distribution for this speciesis therefore evidently, in the main, correct. His citation of the
North American
localitieswhich
have been givenunder
thename pungens
do notshow
the exact distribution. In general, the"pungens" from
northern localities are either pipiens or restuans, thosefrom
points south ofWashington
are quinque- fasciatus',although therangesof the species overlap to.a certain extent. Justhow much we do
notknow.
Specimens from
the moist parts ofNorth
America, the southernAtlantic States to Mississippi Valley,and from
easternMexico
have genitalia of thenormal form
(fig. 1), like those of specimensfrom
theWest
Indies, Guiana,and
Brazil, as well as the Philippinesand
India. In the arid parts of the'United
Statesand
the western coast ofMexico
a modifiedform
of genitalia is seen (fig. 3), representing a distinct race of the species.We
are not acquainted with the line of separation of the forms, as ourmaterial isinsufficient.Our
specimens of the race, forwhich we
proposethename
dipscticus, arefrom
Tndioand
Coachella, Cal. (in the Salton Sink),La
Paz,Baja
Cali- fornia,Acapulco and
Salina Cruz, Mexico.On
the coast of California the specieshasbecome
modified to a specific degree,and we
characterize it hereunder
thename
comitatus (fig. 2).In
Jamaica we
have recognized this species under anothername,
revocator, because the palpi and the proboscis aremore
or less distinctly white-tipped.
The
genitalia arc however, unmodifiedfrom
thenormal
quinquefasciatus form,showing
OF WASHINGTON, VOLUME
XI, 1909 35that the species is but narrowly separated
from
quinquefas- ciatus, if, indeed, it is a true species at all.In the genitalia of the typical
form
(Plate i, fig. 1) the harpes have a small basal projection, larger than in pipiens,but still in a rudimentary condition.
The
harpagones aredividedinto threeplates; thesecond(fig.1, c)is a rather slender plate about as long as the harpes, with
rounded
pointed tip;first plate (fig. 1, d} broad, very long, with bluntly
rounded
tip; third plate (fig. 1,/) essentiallyas in pipiens.
The
species is especially characterizedby
the great length of the central plate of the harpagones,which
is not divided intotwo
portions, as in pipiens.In the genitalia of the race dipseticus (fig. 3), the first
branch of the harpagones isnot especially elongated (fig. 3,c).
The
other characters remain essentially the same, including theflat
and
pointed condition of the second plate (fig. 3, d).Type No.
of the race dipseticus, 12229,U.
S. Nat.Mus.
Culex comitatus, newspecies.
The
genitalia (Plate i, fig. 2) have the general characters of quinquefasciatus, race dipseticus, but differ especially in the character of the second plate of the harpagones (fig. 2, d},which
is no longer a plate, buta-tubular structure, with obliqueopen
tip.The
first plate of the harpagones is broadand
rather long, withrounded
tip, essentially as in dipseticus.The
basal projection of the harpes is very short, but this cannot be espec-ially emphasized, as its apparent condition varies greatly with the position of the mount.
Our
specimens arefrom
National City,San
Diego, Sweet- water Junction,Laguna,
Avalon,Los
Angeles,San
Pedro,San
Luis Obispo,and
Stanford University, California, all these places beingon
the coast, south ofSan
Francisco, oron
the adjacent islands.It is
somewhat
curious that the speciesCulex
quinqucfas- ciatus, afterranging throughout thewarmer
parts of theworld
unchanged, should, in the arid parts of America, developfirst a distinct raceand
finally a species. Thismust
be of significance in regard to the originalhome
of the species. Evidently quin- quefasciatus is of tropicalAmerican
origin,and
has latterly spread,no
doubt through the agency ofcommerce,
to all thewarmer
regions of the world. In these places it has not been resident longenough
to develop local racesand
species, as ithas
done
in America. Conversely, it is probably thatCulex
pipiens is ofEuropean
origin,and
has only latterly spread toAmerica
through the agency ofcommerce,
Culc.v comitatus has the
same
habits as its congener, quin- quefasciatus.The
larvse occur in all sorts of artificialaccum-
ulations of water,and
the adults frequent housesand
attack the inmates at night.Type
No. 12201,U.
S. Nat.Mus.
Culexrestuans Theobald.
The
genitalia (fig. 5) have the harpes with a distinct,though
short hasal projection (fig. 5, &).Harpagones
divided into three plates; first plate (fig. 5, c) narrow, rather short, withrounded
pointed tips; second platebroad, short, the tipbroadly rounded, with small reversed teeth near the middleand
a large, sharp, angular spine projectingoutward from
near the base;third plate (fig. 5, /) essentially as in pipiens, but
shown
fore- shortenedfrom
sideview
in the figure.The
genitalia agree with those of quinquefasciatus in thenumber
of divisions of the harpagones, but are readily distin- guishedby
the peculiarform
of the second division, with itslarge lateral tooth.
This species agrees with the foregoing ones in the vestiture of the
mesonotum, which
consists of the ordinary narrow, curved scales,which
cover the surface rather completely. Inall of the following the vestiture is of minute, hair-like scales, not covering the surface.
The
dentation of second division of the harpagones allies the species with the next group, typifiedby Culex
similis Theobald. This species, therefore, standsby
itself, allied to both the pipiens
and
similis groups, butmore
closely to the former.
Culex salinarius Coquillett.
The
genitalia (Plate in, fig. 7) have the harpes with a long curvedarm from
the outer base (fig. 7, b}.Harpagones
divided intotwo
plates; the outer one (fig. 7, d} a broad plate with a series of long teeth,which
lie nearly in a plane, a large tooth above, partly separated,and
anupcurved
onebelow
are separatedby
arow
of smaller,more
even teeth; inner plate concave, tapering to arounded
tip (fig. 7, /).The
species is especially characterizedby
the absence of the long lateral tooth of the outer plate of the harpagones,which
is present in the
two
following species.This species, together with the following ones, similis
Theo-
bald, factor
Dyar and Knab, and
pro.vimusDyar and Knab, form
a distinct group, separatefrom
the pipiens group.We
shouldnot discuss
them
in this connection, except that the simi- larity of coloration of the adults has led to confusion.The
OF
WASHINGTON,
VOI.UMK XI, 1909 37vestiture of the
mesonotum
is distinctly different in thisgroup from
that of the foregoing f>if>icns group,and
serves readily to distinguish them, without the criterion of themale
genitalia.Culc.r salinarins is a close ally of siniilis Theobald,
and may
be consideredto takeits place in
North
America.Culex
siniilis inhabits the Antilles, while salinarins isfound
throughout the eastern part ofNorth
America,from Maine
to Floridaand westward
tothe Mississippi \alley.The two
species are doubt-less derived
from
acommon American
stock, but have been long separated,and
have developed well-marked specific characters.Cnlc.v factor represents these species along the Pacific coast of .Mexico
and
Central America.Culex factor Dyarand Knab.
In this species,
which we
have not figured, the genitalia are ot the type seen in salinariusand
siniilis.The
outer plate of the harpagones has above a large tooth, followedby
threeslender, well-separated ones below, the lower angle
forming
a largerounded
angle, too largeand
broad to be described as a tooth; a long horn-like tooth arises in a different plane,and
exceeds the other teethin length.The
lowerplateof theharpa- gones is of the usual form.The
harpes have the erect portioncrowned
with'spines as usual, the basal projectionvery longand
curved, as in our figure of salinarius (fig. 7, b}.The
species is especially characterized by the fewness of the central teeth of the outer plate of the harpagones,and
the long horn-like tooth,which
exceeds the other teeth in length.The
scales of the
mesonotum
are hair-likeand
sparse, but they are largerand
coarser than those of salinariusand
siniilis.The
species inhabits the west coast ofMexico and
Central America,seeming
especially athome
in saltwater.Most
of our specimenswere
bred at Salina Cruz, Mexico,from
salt pools behind the beach,and Mr.
Jennings has obtained the species in theLa
Hocaswamp
in the Canal Zone,Panama, which
is a lowswamp
on the Pacific side of the Isthmus.We have
also speci-mens from Tehuantepec
and Santa Lucrecia, Mexico, bredfrom
fresh water,showing
that the species is not confined to salt water. It is evidentlymuch
like our salinarius,which
ismost
abundant
near salt marshes, although generally distributed throughout the country, but perhaps this ismore
addicted to actually salt water than salinarius is.Culexsimilis Theobald.
The
genitalia (Plate in, fig. 8)have
the harpagonesand
harpes essentially as in salinarius, but are especially distin-guished therefrom by the long lateral tooth of the harpagones
(fig. 8, c),
which
arises in a different planefrom
the other teeth. This species ranges throughout the Antilles;we
distin- guish theform
occurringupon
themainland
in theGuianas under
thename
lachrimans. This is probably notmore
than a geographical race, yet the genitalia (fig. 9) apparently differin the shape
and
proportions of the teeth of the harpagones.These
apparently differ in different slides,owing
to the position of the parts,and
toomuch
stress should not be laidupon
these apparent differences. Nevertheless, there areno
doubtsome
actual differences,and
the race is agood
one, aswe
note cer- tain slight differences in the coloration of the adults.The name
lachrimansw
ras proposedby
us (Smiths. Misc.Colls., quart, iss., LIT, 259, 1909) as a substitute for
Culex
aikeniiDyar and Knab
(Proc.U.
S. Nat. Mus.,xxxv,
61, 1908,which was
preoccupiedby Gnophodeomyia
aikeniiAiken
(Brit.Guian.
Med.
Ann., 1906,60, 1907), sinceGnophodeomyia
falls as a
synomym
of Culex.The name Culex
lachrimanswas founded upon
adultsand
larvae. It has since transpired that therewas
a mixture of material, the adults being Culc.v quinquefasciatus, the larvaeCulex
similis.We
have concluded to restrict thename
lachrimans to the larvae,and
it will thus stand for the local race of similis inhabiting the Guianas.Culex proximus, new species.
The
genitalia (Plate u, fig. 6)have
the basal projection of the harpes (fig. 6, b) longand
curved.Harpagones
divided intotwo
plates, the upper one very irregularly shapedand
toothed, a large blunt tooth at the bottom, longand
curved, a similar but shorter one at the top with agroup
of smaller ones between; a long sharp tooth arising in a different planefrom
the others (fig. 6, e)and
exceedingany
ofthem
in length; lower plate concave, broad, withnarrowed rounded
tip (fig.6, /).
The
plate isshown
fully extended in the figureand
appears very broad in comparison withsome
of the other figures, forexample
rcstiians (fig. 5, /), but this difference isdue
to the position of the parts in the slide.The
species isespecially distinguished
by
the length of the lateral tooth of the harpagones (fig. 6, <?),which
exceeds all the other teeth in length.Our
specimens of this speciescome from
the Canal Zone,Panama, and
are in part those referred toby Mr. Busck
asCulex
regulator (Smiths. Misc. Colls., quart, iss., LII, 67, 1908).Culex
regulator is asynonym
ofCulex
similis, towhich
this species is closely allied, butwe
believe that thePHOC.ENT.SOC. WASHINGTON. VOLUMExi,PLATEi
b.
1
C-,
GENITALIA OF MOSQUITOES.
PROC. ENT. SOC.WASHINGTON. VOLUME XI. PLATE
6
GENITALIA OF MOSQUITOES
PROC. ENT. SOC. WASHINGTON. VOLUMEXI, PLATE lit
9
GENITALIA OF MOSQUITOES.
OF WASHINGTON, VOLUME
XI, 1909 39Central
American form
has departed widelyenough from
thecommon
stock to deserve specific distinction.Culex
similis, therefore,exists in theAntilles, gives rise toa race, lachriinans, in the Guianas,and
develops a separate, but closely allied species, proximus, in Central America. This is a parallel development to that of Citle.r quinquefasciatus, referred to above, with its race in the arid regions ofNorth
America, developing a separate speciesupon
the Pacific coast.We have
before us, besides the forgoing, several other species of the salinarins group, but ofwhich we
eitherdo
not possess males or they are so obviously distinct inmarkings
that
we
do not consider it necessary togo
further withthem
in this connection.EXPLANATIONOF PLATES I TO III.
Figures of the basal parts (harpes and harpagones) of the male genitalia of certain species of Culex.
1. Culex quinquefasciatus Say, Iloilo, P. I. (G.
W.
McCoy).2. Culex comitatus Dyar and Knab, Los Angeles, Cal. (Dyar and Caudell).
3. Culex quinquefasciatus variety dipseticus Dyar and Knab, Salina Cruz, Mex. (A. Duges).
4. Culex pipiens Linnaeus, Urbana, 111. (F. Knab).
5. Culex restuans Theobald, West Springfield, Mass. (F. Knab).
6. Culex proximus Dyar and Knab, Taboga I.,
Panama
(A. H.Jennings).
7. Culexsalinarius Coquillett, Chesapeake Beach, Md. (H. G. Dyar).
8. Culexsimilis Theobald, Santo Domingo,
W.
I. (A. Busck).9. Culex similis variety lachrimans Dyar and Knab, Georgetown, British Guiana (E. D. Rowland).
DESCRIPTION OF A NEW MOSQUITO FROM CUBA.
[Diptera, Culicidae.]
BY
HARRISONG.DYAR
and FREDERICK KNAB.Culex
ignobilis,new
species.Proboscis and legs without pale rings; proboscis swollen toward the tip: abdomen without dorsal pale bands, dull blackish, lateral spots yellowish white, basally situated on the segments; venter pale-scaled, with indistinct dark bandstowardthe tip. Occiput with pale scales and erect black forked ones. Scales of the wings broad,
many
obliquely subtruncate.Four
specimens,San Antonio
de los Bafios,Cuba
(J.H.
Pazos).
7 v/>r No. 12239,