l:klkin. Schick Y Heinemann ( 1<:;71 ). La lmica clave t:cncra! cii'p\mibk c" 1a de Lane, 195.' l. la '-'I.

d c'-
L\j'!Crl .~2 e'j1l'cic" en .~el-lln(1" en Aml;rica Central: 7 e"pecies
solo para fnrmal., adultas. Las lan·a" y pupas pu",'dcn "cr
dl' SI/\II/IIUlt entlt' !()'- ('/'ilt/florinl/I'; h l'''pel'lc'' en 1.1I/;·ollliOI)\: y idl'lltitil';lda" ,,\'11\) ha";\ tiihUl'lI c"a cLtve. ~Lly Lllllhll;n Ull.l l·!:t\C
1') npl'Clc" ell Cllf"/'rhrd/il dClltlll dc !u" COJdhll'!lJll;\l'. par;1 ludo' Ill" c..,tadp" P;\I;I .I;\lI\;lll·.I, 1'1l'" . .·IIt.\\h PI" Ih-!klll.
I kllll'll\;\1l1l \' Pa~'",' ( Ilno).

References
Belkin. J.!'J .• R.X. Schick and S. Heinemann. 1968. Lane, J. 19~2. Dixinac e Chaob(lrinac. Revisao dJ . . c"pccie ...
\l\l"quitocs originally described from Aq;entina. Bolivia, neotropicas. Rc\', Entomol. D:X 1-14X.
P,lrJguJY and Urut:uay. Contrib. Amer. Entomol. lnst. 4:<:;-29.
Lane. J. 1953. Ncntrorical Culicidae, Vo\. I. University of S~in
Bdkin. J.l\ .• S.J. Heinemann and W.A. Page. 1970. Culicidae Paulo. Brasil. 54S pr.
Ill' JJlllaica. Cuntrib. Amer. Ent. In,,t. 6:237-2-1.7.
Lane. J. and T.G.H. Aitkin. 1957. Chaobnrinae from Trini(!aJ,
Hdkin, J.H., R.X. Schick and S.J. Heinemann. 1971. B.W.1. Ann. Lntonw\. Soc. Amer. 49:530-5-1.3.
~l(l"quitocs originally described from Brazil. Contrib. Amer.
Miyagi. I. 1975. A hacllIatnphagous Corerhd/a from Jap,\!1.
1'.11\(l11111\. In"t. 7:1·6.1.
JJpart. Jour. Sanitary hlu\. 2():25·2().
Cook. E.E 1956. The Nearctic Chaoborinae (Diptera: Culicidae).
Roth, J.C. and S. Parma. 1970. A Cllllohorus bibiiognphy,
llniv. Minnesota Agric. Exp. Sta. Bull. 21 ~u -102.
Bull. Entomol. Soc. Amn. 16: 100-110,
n~ar. II.G. 1926. i'-:otes of Cort'l/irdla arrendicuiuta Grabham.
Sadhcr. O.A. 1977. Two species of Chuohorus from V.:nezuela.
In.,ee. In"c \knstr. 14:150.
Acta. BiD). \·enCl. 9:IY5-2i2.
D~ar, H.<;. and R.C. Shannon. 192~. The American
Stont'. A. 1969. F:lmily Chauhurid;!\.', fa,l'. J 1: l-S. In: N. P~q'\l':O
Ch.I'l\1llflllac. 1I1\cl.:. Insc. i>1cn"tr. 1~:'::(\1-21fl.
(cd.), A l'atal(,~ul' \1: the l)q1tcra 1'; rh-.' ,\:ll ...·ri...-;, ... I.,(1:J!h (11 tLl
I.anl'. j. 19.39. Nllte" on nl'nhernatl1phagllu" Culicidae. HoI. BioI. United States. Dept. fuo!'. Send. A~ric. S:H) Paulo.
(11-'.) 4:99-1 U.
Williams, .J.A. and J.n. Edman. 1')6~. Oc(urreI1C '':: ()f bi'.",K~
larll'. L 19W_ Nlln hcmat()nha~ou" Culicidae (second paper) Bol. meals in twp srecic~ of Corcr/rrdla in Flnrida. Ann. Ent(l!:1<)I,
In: Hurlbert,S.H.and Villalobos-Figueroa,A.(eds.) 1982: Aquatic
biota of Mexico,Central America and the West Indies.-San
Diego,California,529 pp.

CHIRONOMIDAE
Friedrich Reiss

Introduction
The nematoceran family Chironomidae (syn. Tcndipcdidae) is the depending on the species, the water temperature and tr.~
most important group of aquatic insects both from a qualitative and availability of food for the larvae, The Amazonian srei:icl.,
quantitative point of view, Larvae and pupae of the different Chironomus stren:kei Fittk. requires about 1 week for its life
"peL'ie" colonize all types of natural and artificial waters. such as cycle. \vhcreas Chirol/(ImllS gigas Reiss. a spc;i(.'" inhabiting the
l'uld and hot springs. creeks and river~. Jakes, ponds. small ponds, same region. requires 10-12 weeks. Within thc"e extremes nw"t
small ['nob. axillary waters of plants. as well :1<; rescrv'oirs, water tropical chironomid specics probably produce "cver:_t1 til many
tall\.. '-. canal", and rice fields. Some panheDogcn<:tic species generatiom per year.
(\lIO!1IIl' even urban water supply systems. Ma~"ive occurrences ()f The larval sta~e comprises more than 9(Y; of tl'c life cycle. Pq1;tl
tk !.Inac ~lI1d ima):inc" ofrl:!1 re4uire chcmica!. bil'lot:ica! or and illlat:H1~tl ..,ta~e, an.: . . huI1. The il1l:lgU (l!lly li\c,- fllr 1 In a fl",\
\llll'~'r;lkd ((llltrol r:ll';\'l:n:" i Ci!(',;h~tu" I Ins). f\.hny "pccics of the
Jays. Du:int,: fhi" pL'rllld it fcc,j.., ,)11 pLint l'\.l!lbt~',. ;!nd tlLI' tll.:
,ubl.llllily Ortrh1l'Ltdllna<.: kad "crnircrrc<.;t!lal iin:s in nHli"t cnl'f~y IC\juirl'lII""llh (1f rl';)rt)dll(ti\ll1 ;~r ...' 1J1~'! C\'i'til.lii,'il ~\"U;l:l\
h.lb!!.I!' or arL' c(l[npktc!y tl'rre,tll:iI. hlfthcrl1lore. the take, place 111 till' ;111 Wlll'll sillrk kr'Llk.., .Il ' i';(l,l,·h :1;<.' ,,\\.;[!lii;:~·
('I!II'lll(lIllH\;t,,: i" th.: Ilfl!y 1Jl'.n:t Ltnlilv til:lt II:!" c"lolll/cd the
11I.lle ... S\lllt.: "llL'l'lL" ;1I\.' kllllWll t(l ,'''FuLlll' OJ) ,I "ld,,!!.l!,'. '.t!,li .: ...
1I1.1111ll· liltllr.d hl'lifh"", and I" rl"prl"l'IIl<'d [i1lrc by ,n'cr:tl t!L'lIl'1<l
tht: "0", \';tin. \11 \q'CLltlllll
.1I:d "'Pl'll\, ... III the subtaillihl..'" 'kllJlatu~'ct(lnill;le, Or1hm:ladllluc
alld (·illnlllol1lin:tc. Thl' larvae of mall}' ci1lfolloll1iJ ",pL·lie . . SilllW h,l!>l!.lt prl'!"!~'ll\·t",.
They can therefore he lIsed a" indic"tnr or~~l!1!'im" for \\ :\\L'r
Chltl'1l11rniJ larvac arc mainly herbivores or detritovon:s. Some quality and ih altcmation by m~ll1\ activities.
1,1\<1 ;Ul' partially or obli,l2atorily carnivorous. e.g. species of the
.. uht.mllly 'bnyp\)dinae and of the Ifamischia group of genera in Chironornidae occur worldwide. Many specics ha\'l' wide
the "uhtamily Chironl)minae geographic di ... trihutions \.\'hich, hll eX;i'npk, m:ly compri"c tbe
entire H\)!arctic or Palac\1tropic rl'!,;ion .... Only a few Uxa ha\c
The {(ltll 1Ill't<llT1orphosi-;. from egg to the hatched imago. requires been found to bl..' restricteJ to smal! gl'o12raphic areas; e. ~. s<'me
11\1111 a minimum of I weck to a maximum of several years,
species of the Pod()nl)minal' are cfll.kmic to the southern And.: ....

4J3
lTp tonow no "pecic'> of chironomid is known to he endemic to a Totally unknown is the hi~h mountain fauna of Centr;1I Am,·!h.•I. II
"in~k river sy~tcm or mountain ran~e. Speciation seems to is unknown \\ hether clement." llf the Nearctic d1irllnolllld I.lllll.1 .Ht"
pr;)ceeJ \ery slowly in the Chironomidae. prohably hecause of prc'>ent in this type of envirunment a!>. thL'Y are Oil till' \\lk ,11111\ \11
thl'ir ~reat capahilities for dispersal. Pkistocene-originated the highlands of Central \kxico (. . ee RL'i,,~ I l02a. Sell;! lll'lll
~r\xl;l!i()I1" a . . /t)ulld in other ~rnups of aquatic insects. are not yet 1977b).
knm\ n for the chirlHh)mids.
Some marine chironomid "pL'ciL'''i he1nnging to the "'UOLlIlldlL'\
At prl·\L'nl. nnly [()ll~h c-.tirnates of the numher of recent species Telmatogetoninae (Wirth 19(1»). Orthocladiin:lL' (l{(lr;I\.·j.; 11171 h)
can he made. In Lurnpe. where the chinHlOmid fauna is known and Chirnnnmin;te (BrehChko Il)X I) arL' kn()\\. n Irom thl' ('clltr ,II
hctter than anywhcre cl"e in the world. 1..+04 "pecics have heen A me ric a n coa"t. The mo" t rece n t com pJl at illn on Ill;tr I Ill'
rq'i..,kred. Out of thc"e, 9.1~; have heen de.;cnhed within a period chironomids including a key to the genaa i~ Ha"himntll ( 197h).
of on I y I () YL' ar" (Fi tt kau and Rc i ss 197 Xl. H O\l,'e \"Cr, the For the ncar future. studi" . . cnncerning chironomid"i of CL'ntr;,1
chirnll()lllid fauna In lar),!e parts of Europe j" imufficiently known America should he primarily faunistic and taxonomic In
or Iwt krwwll at all. Thi" i" true tor the ~1cditerranean re~ion and orientation. Only through this approach can a !>.olid foundation for
the tundra of 11llrti1crl1 Europe. It i" to he su"pected that subsequent ecological and applied "tudics be laid,
6.()On 10.(\(1(1 l'himn,)fl1id "pl'l'ie.., e'{j..,t wnrldv, ide.

The t,lllll\\ il1~ paplT . . can he recommended as an introduction to

thl' !;I'(1Tlll!ll~ ;ll1d It:rlnin{)I()~~: of thL' Chironolllitiae: Brundin
11)'\lla. Il)(,/): Ftttkau I<)h2: Hirh'noja 11)73: Rphack 11)71a.
II»/l.l. 1\)77h. 1975a: S,Il'thcr )lnlh. 1l)77c, 19X(): \LI . . (ln 191.'. La familia nl'mall)L'era de \(l\ Chirnl1(lmidac (<'If1. 'Ii:ndipedilbe) l'"
J\dd!!lIl'lal rl·t'I'fL'fll'l'\ may Ol' found in the chirpnllmid el ~~rllp() m:;.., imp(llL!Il!L' lk in\el:!ll., acu.iti~·ll"', tanto dC\~k cI
bd':lt l ,:i'If'll\ III httl-...iLi, Rl·I.., ..... ;!;]d l1utlrILhkr (llJ7()) and in Ih pUlltll dL' \ 1,,(<1 L'llantlt;lIi\ () l'om\l L·ualIL,tIVO. 1.:(\ Lifva, y pup;" tiL'
~U!)pkrIlL'nt (1llllfrichtcr and Rei~, I l)X I). J-urthermore. a la . . di'L'r\a, L'~p(:t'il''; Collllli/an !utln tl[1U de at'll:!. tanto n:tturak,
pUll lllil'al l·,i"r". "Chlwnu!1lu" - i\L\\~ktter on Chiro,wrnid como artlficlaiL'-;, tak . . C()fIIU t IIl'nle" fria" y tL'rlll;dl'';. 4uehrada-;,
Rc"carL'i1" (httbu and Rei"s 1')67-1976, Fittbu. Rei,,, and bgm. l.·har,:as, [X'quelHh dWfCO-., "t!ua axilar de Lt . . plan!;! .... a"i
Subktt . . 1<)7Xn.1. \\ hich serves as an up-to-date information como re-'L'rvorios. tanqllc,; de agua. canale . . y (ampo~ de alTol.
~()Url'C In l·hiwnomidolo!!y. AI),!unas c . . pecics par!t.:rHlgl:nc!icas colol1ilan :tlJl1 los \1 ... tL'rna.; dL'
The bihll\l~raphy of R.()~enberg (11)7S) li,>ts a selection of agua urhan,)s. La occlIIl'nl'ia ma . . iva de larvas e imago . . requi,'re a
impllrtant p.lper" on <'Jmpling methods and sample analysis for the menudo tratamientlls quimico". hillklglCOS 0 integrado" (Grodhaus
m;lcn)/oobenthos. 1975). Muchas espl'cie~ de la suhfamilia Orthoc1adiinac "on
semiteITes!res en hjhital" hunh:d~)s 0 compklamentc terre . . trcs,
The ci1ironllll1ids of Central America arc much le,>s well known Mas aun. Il)'; Chironilinidac <,(In la unica familia de insecto.., que ha
than thp . . e of trnpical SOllth America. So far only ahout XO "pel'ies colnllilado el bellto-. IItmal marino y e,tan rcpre.;cntados alii por
han' ht'cn identlfll'd. However. it can he rnul2hly e-.timated that the varins rt~nems y C ... I"X,cie . . de las suhfarnilias TcJmatogetoninae.
I1Ulllnn pf "Pl'I.·!'-" ..,hlluld he 1)00 - 2000. A great many of these Ortho(laLiiinac y ChirolHlll1lnac.
"pccic~ probahly will also occur on the South American continent.
There i.; no Central American lake or river in \\hich the Las larvas de los quiron6midos son principal mente herhivoras 0
chircnlllniJ fauna has been "atisfactnrily analysed qualitatively or detritivoras, Algunos taxa son carnfvoros parcialcs u ohligados. p.
quantitatively. Therefore. any generalizations concerning the cj .. especics de Ia suhLtmilia Tanypodinae y del grupo Hamischill
chirlllHl'nid fauna of Central America must be considered de gcncros en la subfamilia Chironomin:.J.c.
tentative. La metamorf6sis total. de "de el huevo hasta cl imago ec1osilmadl).
Central America helongs to the Neotropical faunal region. with the requiere un minimo dc una semana hasta un maximo de varios
exception of most of Mexico. and its chironomid fauna is anos. <.kpendicndo de la cspecie, de !J temperatura del agua y de la
Q,11du[('t\ accordin),!ly. Only a fcv, specie" appear to have their di"ponihilidad de alimento para las larvas. La c!-.pecic amatlmica
orl~in in the Nearl'lic region. LikC\\i . . e. only a few of the
Chir(lf/Om:lS srrclI:kt'i httk. fl'ljuiere c(rca de una scmana para su
\clllrop:cd t;l \a "(,L'm to h:lv(' mi~r;ltcd to the ~can:tic region. and cicio de vida. 1111(:nt::1" qllL Clzirollo/!1uS Xigus Retss, una C"I'X:cic
tho"l' r:l(l'.t\ pnly :I" I:tr ;1 .... the <.,putherTl purtioll. A typical clement que hahit~l ]a Illi"ma rq!i(ln. nl·ce . . ita 10 a 12 semanas. La mayurla
of (he :\C('l!opi,..:al fauila i" the )2enu\ Go,-/ciichiroIlOfllUS, which is de las e'.pecic" trllpi\.. ak . . de Chiwnomidae producen varia,> ()
r~'i)rneflll'd hy '.ner;!1 ~recie.; iii Ccntral America. hut which is
rnucha.., gclleraclone . . por aI'1\), dentm de los extrel110s scrlalados.
a~'l) prc'~'llt In the . . illjth"TIl l\1111'd Stdll'-.. Los ("ladlo\ !an ;tk'i ab:uL"trI j;)js del 9()(i( del C1ch) vital. Los
e~tadios de pupa e lIll:l!C(\ .... (In cllrtll..,. EI ima)...:o vive un dia 0 pocos
Till' cndl'IllIClty Ill' chlflllhlrnid-. in (\'l1tral America j.; difficult to
dia~. Durante (,,,Ie peril 1dl) . . 1.' ;.dl!l1enta de las e'{udaciones dL' las
CV;t!U;IIC at p!e~,ent. Sc\eral l'hir,lI1omid )2cnera and specie.; have
been idl.'l1:ifil·d that so far h.lve occn found only' in Central planta'i. cumpliendo as! COil 1o" rcqucrimicnto-. cnergcric()s para la
America. H(lwever. their full ),!eo!,!raphic di-.tribution is not reproducci,\n. La copula generalmcnte ticnc lugar en el aire
known. The ell.'\atilln~d variation of the Central American cuanLlo una hemhra se aCCfC:l al enjambre dL' machos. Algunas
e~pccies copulan en su~[r~!tos talcs como cl sudo, la supaficic del
chir,H1t)m:d Luna is cUITL'ntly hard tn delermine. In the l(mlands
we tind pred,l[11I11antly ~pecics of the suhfamilies Chironominae agua y la vegctacll1n.
;ind TmYP(ldin;\e, as is usually the case in the tropics. In the Las larvas dc muchas especies de quirnn6midos mucstran una
rUlIllin)2 water . . of the mountainous region". species of the fuertc sL'Jcctividad por los h.lhitats, Por 10 tanto, puedcn ser usados
subfamily Orthoc1adilllae "cern In he more ahundant. There are como organi"mo" indicadl)rc~ de la caliJad del agua y de las
indil·ation . . th,lt "(ll11e ,t:cnera of the . . ubfamily Orthoc1adiinae may alteraciones que clla ..,ufrc pur la actividad humana.
h(' endellli,-' to Central America and may hc related to African taxa
Los ChirnI10midac c'{i . . !cn en totlo cI mundo. Muchas especies
(Sactlier 19x I). \1oulltain streams of Central America seem to
tienen distribucionL's gcograficas amplias. las cualcs pueden
COllf;lin more repre"entative<, of the '>uhfamily Orthocladiinac than
comprender. por ejemplo, toda la regi6n Hol~irtica. 0 e1
C(lI11I';ILlhk \trL':1Il1" PI' tfopil'al South Amt:rica. Thcrefore, Central
Pa1co[f()pico. Solo unos pocm taxa ticnL'n areas geogr:'ificas
AlllCril';l pU\"lbly l'Oldd he viewed a . . a /oorl'ographically
IX'quclla~: p. ej., alplIla" e"pecics de Podonominac son cndemicos
illlkpclldent suhrq.:iol1 ..... eparate fmm tropical South America.

434
de 10 ... Andes surerlOs. lla...ta ahora no se Cllnoce ningun quirnn6midos c"ui cstrul:turada de manna corre<,pondicl1tL'. St"10
qllirPIH1mido quc ... ea endcmico de lin solo sistema hidrogrMico 0 una ... pocas especies parecen hahcrse ori~inado en la rq!llin
C\lkna montaiiosa. La espceiaeion parcel' scr lenta cn los Neartica. Por otra parte. s610 un numcro redu.:ido dc taxa
Chlfll(wmidae. dchdo probablcmentc a su gran capJcidad de neotropicales parcce habcrse extendido a la region Nea;tica y estos
dl\l'cr~i{m. La especiaci()n de orfgcn p1cistocenica que se en su mayorfa. lIegan s610 haqa la parte sur de esta region. Un
l'1ll'UClltL.l en o:ro" grupos de insectos acu;iticos, no se conoce en elemento tfpico de la fauna neotropical es cJ gencro
111\ quirl1!H)lllidos. Gocldlchironomlls. el cual esta reprc'ocn~ado pm varias e:-pccics en
A,:tll.t1mcntc st')lo ~e puedcn haccr torpcs estima('iones del numero America Central pem tambien sc encucntra en cl sur de k)s Esudus
de C'f''l'I(' recientes. En Europa. dO;1JC Ia fauna de quiron6midos
Unidos.
'c \.'1)11\1\.'( Ill\.',l()r que en partc algu:la, sc han rcgistrado I A04 Acrualmentc cJ endcmismo de los quiron6mido... en Am~nl.:a
c'pccics lk cstas. 9. I (;f, se han dcscrito en un pc dodo de Central e ... dificil de e\·aluar. Han sido idcntificados varios gcncros
W!.lllll'nll' J() ~1I10S (Fittkau y Rci ... s 197X). Sin embargo. la fauna y especics de quiron()midos que hasta la fccha s('llo ... " han
Ill' qU!rI)IHimld()" csta in ... uficientemcnte conncida en muchas encontrado en America Ccntral. Sin emhargn. su dl ... trihucil.l!1
p.lIte, dt: Lumra. 0 es cnmpktalllcnte dc ... conocida. A... f IK'l)rrC en gcogr{ifica completa. se licscono('c aun. Las vari~H:ioncs
1.1 rl'~!\ln dd Ilh.:ditcrr.inco y en la tundra del norte de l::urllpa. Sc altitudinalcs de la fauna de quiron()mid()s ccntftlaml.'ric:ltw ... ,on
pucdc 'ompcchar que en eI mundo existen 6.000 a 10.000 especies dificilcs de dcterminar. En las tierra ... bajas encontramn ... Ci,H1lO
de ljlllfllJ1l11nidos. predominantes a espccics de las subfamilias Chironominae y
\'1\, \I~uit'ntc ... trabajos puc den recomcndarse como introducci6n a Tanypodinae. como suck ser el caso cn las fl'giones tropicalcs. En
1.1 1.IWIlOlllia y a la terminologfa de los Chironomidae: Brundin aguas corrientes de las rq:il.mcs montail{)"as, p.lrCl·cn ser mas
111<ih,l. Jl)66; Fittkau 1962; Hirvcnoja 1973; Rohack 1971a. abundantes especies de Ia !o.ubfamil ia Orthlh:ladi mac. Hay
l'i7I',I. 1l)77~. 19na: Saether 1971h. 1977c. 19S0; \bson 1973. indicaciones que algunos gcneros dc la suhfamilia Orthodadlinae
I'tH'den halLtr',e rcfercncia" adicic na1cs en la hihli(lgrafia de
1
puedcn ser cndemicos en America Central y pudie"en e"tar
'lilli"!l\)11lid\l\, de Fittkau. Rei" ... y Hnffri~'hter (11)76) Y cn relacionados con taxa african!~s (Saethcr I ()S I\. Lo'o arroyns de
\!:;,L,·Pll.'flt!l i Huffri,'htc: y Rei"" 1l)~ 1)' \Li" aun. existe una n1t'ntana de America Central parccen conte ncr mas representantes
l'td'lic,Il'llll1 pl·r;\)dll'C!. "Chinll1t1I11Us ,- t\cw:--tcttcr on Chin'nnrnid dc la suhfami:ia OrtbncLtdiin,!c que arruytls <,cmc;antc;;, de
Rc'c.lrl.·h" (hUkau y Reiss 19(,7-1476. Fittkau. Rei ... " y Suhlette Sudamerica twplcal. Pur 10 tanto. po"iblemcl1tc se podria
1'J'''S ~ '!l' ) qu;: ... irvc cumo fuente de inf(\rrna(:i~')n al.:tu;.t1ilada en cOI1siderar a America Central como una subrcgil'lO loogC()gr{lfica
qUIII'IH'lIl1d\)!0t: ia
di ... tinta de Sudamcrica tropil"11.

I .1 bll'll\)~raria
dc Rosenherg (19 7 8) li"ta una seleccion de trahajos La fauna de alta mnntana dc America Central es totalmente
IIIlP,l(t;lllte'o sobrc metodos y analisis de mucstreo para el desconocida. No se sahe si estan prcsentes ekmentos de la fauna
11)," n~/\l"t~ntl)S.
de quironomidos ncarticns como In estan en los vokancs de las
tierras altas de Mexico central (vcr: Reiss 1972a. Serra-Tosio
I '" ljlllwn6midos de America Central son mucho mcnos 1977b).
\III]I)o,.hll)'" qlle :.:qucllos de Sudamerica tropical. S('!amente 80
npnll'\ h,lI1 "ido ientitil'adas ha"ta la fecha. Sin embargo se puede En las costas de America Central ~e conocen algunas cspecies de
("tl11\.1[ .l'11l"cramente quc el ntlIllerO de espccies serla entre 1500 y quiron6midos marinos pertenecientes a las subfamilias
~(HI(I I!n .l'ran numcro de estas especies probahlcmente se
Tclmatogetoniinae (Wirth 1969), Orthocladiinae (Roback 197 J b)
('1";.Jl'lltr.1 tamhicn en Sudamerica. No hay ningun lago ni rfo Y Chironominac (Bretschko 19X I). La rccopilacic'ln mas rCl:lcl1te
U'lllrll;lllll'flcano para cl cual la fauna de quironomidos haya sido de los quironomidos marinos ha sido rcalilaJa por Ha ... himoto
, .1 I I , I a t' II) r I a men tea n a liz a d a. y a sea c II a lit a t i v a 0
(1976) e incluyc una clavc para los gCI1eros.
l'U.llllll.ltIV;I111cntc. POf 10 tanto. cualquier gcneralizacion Para el futuro cercano. los e"tudios sohre los quironl)midos de
\.IHkl'lIllcllte a la fauna de quiron()midos de Am¢rica Ccntral debe America Central dcberan tener una orientacion primariaml'nte
..cr ..:pn"'llh.'rada tcntativa. faunlstica y taxon6mica. Solamente de esta manera se comtruira
Con Ia C'l'.cqx:i()n de la mayor parte de Mexico. America Central una base s6lido para la realizaci6n de posteriores estudios
r~.'rtrnccc a la region faunhtica Neutropical y su fauna de cco16gicos y aplicados.

References
~\lt(': The letter designations accompanying the year of (Diptera. Chironomidac) in the Caribbean Sea. Smithsonian
l'uhl1l'.ltioll tor certain citations cprrcspond in all cases to the letter Contr. Mar. Sci. (in prcss). Larvae and female pupac of a prohahly
d"I)':l,!II""'" elllployed in the "ihlio~·raphic-. of Fittkau. Reiss and partheno!!cnetic srcl'ie~ of Pontonlyia twm the cpast of lklile.
1i'::II,lttl'f \)')7(,) and Hul1rid1tcr and Rei ...... (llJXl). Larval' f()und in snillllclIts in waIn as del'p a'> ,~() m,
Bn.'lOnik. P.L. and J.L. Fox. 1'>7 .... The 11Il1llllll'gy llf "l'kl'ted
n"~. EX. 1'>6~ h. An an~!Iy"is of the "sayulc" (Diptera:
Guatemalan lakes. Hydrohiologia 45: 467-4~7. Some quantit:ttive
n u i ~ a n c cat San Carl () s. N i car a g u a. and
l h I I " 11 \ \ III \\ L! I')
estimatcs of larval ahundances of different chirm1l'mid t!.::n..:ra
ICl.\ll1lll1l'lIlbtIlHl\ for isis alleVIation. World Health Organization
(Coc/ot(1nYPlis. Procladius. Chirollomus. Einfcldia; in the
I II\, ,,,'fll'''' W HOE B L20. WHOVector ControlS6; 18 pp.
benthos.
\ rllilll('''f.'L)
Brundin, 1... 1956a. Zur Systcmatik der OrthocJadiinae (Dipt.,
nu~. L(, .• L Dominguez and N. Escohar. 1966. A second
Chironomidae). Rep. lnst. Frcshwat. Res. Drottningholm 37:
qll\l'~ ld' rhl' l'hironomid midge problem at San Carlos.
5-185.
i\" .1' .1):11.1 \\'\'r1d lkalth Or~ani/ation EBL series WHOEBL66.
;,~ I) PI'. (1llI!llC(l~r.) Brundin, L. 1966. Transantarctic relationships and their
sign i ficance. ev idenced by charonomid midges. With a
Un'hrhko. (;. l'>SI. First record of the genus POf1(omyia Edw.
monograph of the subfamilies Podonominac and Aphrotcniinae

435
and the austral Heptagyiae. K. Svenska Vetensk. Akad. Hand!.. I I : (Chironomidae). Psyche (Camhridge) 39: 69-71. Iksmptllll\ \11

1-.+72 . the male imago of Chirollomus pseudojascillta n.sp.

Cheng, L. and C.L. Ho~ue. 1974. New distribution and habitat Gerry, D.1. 1933. Four new species of Chironomidac frlllll Ihl'
records of hiting midges and mangrove flies from the coast of Greater Antilles. Psyche (Cambridge) '+0: 9'+-97. Dcscnpllllll III
southern Baja California. Mexico (Diptera: Ceratopogonidac. the male imago of a new species and the male and femall- 1I11;lI-'0\
Culicidae. Chironomidae. and Phoridae). Ent. News 85: 211-218. of two new Chironmnlls species.
Report of Smittia sp.
Goulden, C .E. 1966b. The animal microfossils of La~lIn;l de
Cockerell, T.n.A. 1915. A new midge from Guatemala. Can. Petenxil. Guatemala. Mem. Connecticut Acad. Arh Sl..'i. '7:
Ent . .+7: 315-316. Description of Chironomus guaitemllitecus 84-120.
n.sp. 9.
Grodhaus, G. 1975. Bibliography of chironomid midge nui\;lIl~'C:
Coquillett, D. W. 1900b. Report on a collection of dipterous and control. California Vector Views 22:71-R I.
insects from Puerto Rico. Proc. U.S. Natn. Mus. 22: 249-270.
Harrison, A.D. and J.J. Rankin. 1975. Forest litter and "Ul';1111
Cowgill, li.M. and G.E. Hutchinson. 1966. La Aguada de Santa fauna on a tropical island, St. Vincent. West Indies. Verh. 1111
Ana Vieja: The history of a pond in Guatemala. Arch. Hydrohiol. Verein. Limno\. 19: 17'36-1745. Larval ahundan~:c of some
62: .:n5-372. chironomid species in tropical mountain hrooks and strl'ams.
Curran, C.H. 1928. Insects of Porto Rico and the Virgin Islands. Harrison, A.D. and J.J. Rankin. 1976. Hydrohinlo!,!ical .. tudIC\
Diptera or two-winged flies. In: Scientific survey of Porto Rico of Eastern Lesser Antillean Islands. I. St. Vincent: Freshw;lkr
and the Virgin blands. (New )hrk Academy of Sciences) I I: hahitats and water chemistry. Arch. Hydrobio1. Suppl. 50: 96-144.
10- 13 Description of two new Cricotopus species from Puerto Ecological data on 24 chironomid species from various standing
Rico and Cuha. and t10wing waters at different elevations.
Curl1', L.L. 1974. The midge fauna (Diptera. Chironomidae) of Hashimoto, II. 1976. Non-biting midges of marine hahit:t"
the L:. S. and British Virgin Islands; the genera Goeldich/rmlOmus (Diptcra: Chironomidae), pp. 377-414. In: L. Ch~n~ (cd.). Mafln~
and S/o/illlyill. Ent. Tidskr. 95 (Suppl.): 58-65. Description of the inserts. North Holland Pub\. Co .. Amsterdam, Oxford, Ne ....
complete metamorphosis of G. ama;:onicus and G. /toioprasillus. )hrk. Summary of all known species of marine chironolllid .. ;
with a hricf statement on biology and distrihution in the Virgin includes a key to genera.
Islands.
lIinenojn, M. 1973. Revision dcr Gattung Cricotl'{1IH van dl'r
Curry, L.L. and M.C. Curry. 1971. The Chironomioae (Diptera) Wulp und ihrcr Vcrwandten (Diptera. Chironomidae). Ann. Zoo!.
found in the Virgin Islands and Anegada. British Virgin Islands. Fenn. 10: 1-363.
Can. Ent. 103: 310-314. An annotated list of 16 chironomid
Hoffrichter, O. and F. Reiss. 1980. Supplement I. A
species from 12 genera; based on larvae and partially reared
bibliography of the Chironomidac. Gunncria 37:68 pp. Lists ahout
imagos.
1800 additional works on chironomids puhlishcd hcfore mid-1979.
Elliott, J.M. and P.A. Thllett. 1978. A bibliography of samplers
Hynes, II.B.N. 1971. Zonation of the invertebrate fauna in a West
for henthic invertebrates. Occas. PubIs. Freshwat. BioI. Assoc.
Indian stream. Hydrobiologia 38: 1-8. Vertical zonation of
(Amhlcside) 4: 61 pp.
Tanytarsini larval abundance in a Trinidad stream.
Fittkau, E.J. 1962. Die Tanypodinae (Diptera, Chironomidae).
Johannsen, O.A. 1938. New species of Nematocera from Puerto
Die Trihus Anatopyniini. Macropelopiini und Pentaneurini. Abh.
Rico. J. Agric. Univ. Puerto Rico 22: 219-226. New descriptions
Larvalsyst. Insekten 6: 1-453.
of the imagos of 5 chironomid species. with mention of some other
Fittkau, 1968b. Sio/imyill amazonica n. gen. n. spec .. eine species.
flllgtahigc Chironol11ide (Diptera) mit einem Hypopygium
Johnson, C.\V. 1913. The dipteran fauna of Bermuda. Ann. Em.
invcrsllm. Amazoniana 1: 259-265. First record from Nicaragua.
Soc. Amer. 6: 443-452.ChirorlOmus cristatus Fabr.. Orrhoc/adiuJ
Fittkau, E.J. 1978. 5ich ahzeichnende Verbreitungsmuster in der sp. and Afetriocnemus knabi Coq. arc recorded.
neotropischen-nearktischcn Chironomidenfauna. Mitt. Deut. Ges.
Kieffer, J.J. 1921c. Description de quelques Chironomides
All!!. Angcw. En!. 1: 77-'1) I. General zoogeographic aspect of
exotiqucs. AnnIs Soc. Scient. Bruxellcs 40: 181-186. Description
wa;m- and cold-adapted chironomid taxa in the Neotropical
of male imago of Nilodonan hctcmchirus n.sp. from Trinidad.
region.
Laessle, A.M. 1961. A micro-limnological study of Jamaican
Fittkau, E.J. and E Reiss, (cds.). 1967 - 1976. Chironomus.
bromeliads. Ecology 42: 490-517. Reports larvae of Pentaneura
Mitteilungen aus lier Chirnnomidenkunde. PIon.
sp .. Chironomus sp. and Cry[>tochirollomus sp.
Fittkau, E.J. and E Reiss. 1979. Die zoogeographische
Lichtenberg, R. 1979b. Chironomiden aus Kolumhien I. Ein
Sonderstc1lung der ncotropischen Chironomiden (Diptera).
Beitrag zur Verbreitung von Cryptoc!adopl'lma hoydi (Beck.
Spixiana 2: 273-2RO. Summarizes present systematic and
1972) (Diptera, Nematocera). Annln Naturh. Mus. Wien 82:
zoogeographic understanding of the Chironomidae of tropical
541-548. First report of Ciadopeima boydi from Mexico.
South American lowlands. A still largely unknown part of this
fauna is to he expected in Middle America. Locw, II. 186lb. Diptera aliquot in insula Cuha collecta. Wien.
Ent. Mschr. 5: 33-43. Description of male and female imagos of
Fittkau, E.J., F. Reiss ami O. HofTrichter. 1976. A bibliography
Chirvnomus octopUllctalus n.sp.
of the Chironomidae. Gunneria 26: 177 pp. Lists about 6400
works. puhlished up through 1974, which deal entirely or in part Miller, A.C. 1971. Observations on the Chironomidae (Diptera)
with chironomids. inhabiting the Ieafaxils of two species of Bromcliaccae on 5t.
John. U.S. Virgin Islands. Can. Ent. 103: 391-396. Abundance of
.'ittkau, E.J., F. Reiss and J.E. Sublette (eds.). 1978ff.
larvae of Tanytarsus sp .• Metrionemus sp. and Penlaneura sp.
Chironomus. Mitteilungen aus der Chironomidenkunde.
MUnchen. Pearse, A.S. 1936. Chironomid larvae from Yucatan. p. 151. In:
A.S. Pearse, E.P. Creaser and EG. Hall, The cenotes of Yucatan.
Gerry, D.1. 1932. A new species of Chironomus from Jamaica

436
A zoological and hydrological survey. Carnegie lnst. Washington. Roback, S.S. 1976a. The immature chironomids of the Eastern
304 pp. Records the larva or imago of three inadequately United States: I. Introduction and Tanypodinae - Coclotanypodini.
dClcnnincd chironomid species. Proc. Acad. Nat. Sci. Philadelphia 127: 147-201. Major work on
the taxonomy of New World tanypodincs; metamorphoses;
I't'arse, A.S. 1938. In~ects from Yucatan caves. pp. 237. 244. In:
A.S. Pear<,e (cd.). Fauna of the caves of Yucatan. Carnegie Inst..
identification keys.
\l,'a,hinglon. Records the pupae of Chironomus sp. and Roback, S.S. 1977b. The immature chironomids of the Eastern
I'.I/·w!ochironomus sp. United States: II. Tanypodinae - Tanypodini. Proc. Acad. Nat. Sci.
Picado, C. 1913. Les bromeIiacees epiphytes consideres comme Philadelphia 128: 55-88.
nubeu bio!ogique. Bull. Scient. Fr. Bclg .. 7'= Ser. 47: 280-295. Roback, S.S. 1978a. The immature chironomids of the Eastern
Good descriptions of the complete metamorphoses of United States: III. Tanypodinae - Anatopyniini. Macropclopiini
"Ahlahesmyia" costarricensis n. sp .• Metriocllemus and Natarsiini. Proc. Acad. Nat. Sci. Philadelphia 129: 151-202.
(lndominojh.l\·atus n. sp .. Cltiroc/adius pedipalpus n. gen .. n. sp. Roback. S.S. and K.J. Tcnncssen. 1978. The immature stages of
and Chironomus sp. from Costa Rica; includes some information Dja/mahatis[a pulcher (= Procladius (Ca!otaIlYpus) pulcher
on ecology. (Joh.». Proc. Acad. Nat. Sci. Philadelphia 130: 11-20. First
Pocy, E 1851. Memorias sobre la historia natural de la Isla de reliable report of the genus Djalmabatista. previously regarded as
Cuba, acompanadas de sumarios latinos y extractos en francees. endemic to the Ncotropical region. from the Nearctic. Procladius
Vol. I. 463 pp. Habana. (Ca!otan_\'pus) Roback. 1971 is a nov. syn. of Dja!maharista.
Rdss, E 1972a. Die Tanytarsini (Chironomidae. Diptera) Rosenberg. D.M. 1978. Practical sampling of freshwater
Sudd1i1es und Westpatagoniens. Mit Hinweisen auf die macrozoobenthos: A bibliography of useful texts. reviews. and
l;II1Ylar .. ini-Fauna der Neotropis. Stud. Neotrop. Fauna 7: 49-94. recent papers. Tech. Rep. Can. Fish. Mar. Servo 7l)O: 15 pp.
[)c\l'ription of the male imago of Paratanycarsus rolucensis n. sp Saelher. O.A. 1971b. Notes on general morphology and
twm Mexico.
terminology of the Chironomidae (Diptera), Can. Ent. 103:
Reiss, E 1974c. Die in stehenden Gewa<,sern der Neotropis 1237-1260.
vcrhreilcte Chironomidengattung Goe/dichironomus Fittkau
Saelher, O.A. 1977c. Female genitalia in Chironomidae and other
(Illptcra. Imecta). Stud. Neotrop. Fauna 9: 95-122. Sio/imyia as
Ncmatocera: Morphology. phylogenie~. keys. Bull. Fish. Res. Bd.
nov ... yn. of Goeldichironomus. New record of C. ama:onicus in
Canada ·197: 209 pp.
l'uha and Mexico. and of G. ho!oprasinus in Cuba. First report of
G JluctudflJ n. sp. from the Virgin Islands. Saether, O.A. 1980a. A glossary of chironomid morphology
terminology (Diptera: Chironomidae). Ent. Scand. Supp!. 14:
Rohack, S.S. 1962a. Some new Tendipedidae from the Canal 1-51.
Z~lnC. Acad. Nat. Sci. Philadelphia. Norulae Naturae 355: 1-10.
lk,~'riptions of the imagos of Thienenzaniella spreta n. sp .• Saether, O.A. 1981. Orthocladiinae (Chironomidae: Diptera)
C"CtltO{Jll.\ tanis n. sp .. C. oris n. sp .. Cantom. . . ia cara n. gen. n. from the British West Indies. with descriptions of Antillocladius n.
'po and Chirunomus aversa n. sp .. as well as further data on gen., Lipuromerriocnemus n. gen .• Compterosmittia n. gen,. and
Je'l'nbed 'pccies. Identification key for neotropical Cricorvpus Dip!osmittia n. gen. Ent. Scand. Suppl. 16:46 pp. Description of
'1'C(le, (male and female imagos). 12 new species (male and. in most cases. female imagos; juvenile
stages in some species) from mountain brooks of St. Vincent and
Rohack, S.S. 1963c. New neotropical Coelo[anypus (Diptera.
St. Lucia.
'IC:ndipt·Jidae. Pe1opiinae). Ent. News 74: 169-176. Description of
the male imago of Coelotanypus cleris n. sp. from Jamaica. Schlee, D. and W. Glockner. 1978. Bernstein. Bernsteine und
Bernstein-Fossilien. Stuttg. Beitr. Naturk. Ser. C. 8: 72 pp. First
Rohack. S.S. 1964a. A new Pelopia from Mexico (Diptera:
report of fossil Chironominae and Orthocladiinae from the amber
TcnJipcdldae). Ent. News 75: 141-143. Description of the male
of the Oligocene formation of the Dominican Republic.
Im;lgo of nmypus catemaco n. sp.
Serra-Tosio, B. 1977b. Deux nouvelles especes de Diamesinac du
Rohack, S.S. 1964b. New record of Coelotanypus cleric Roback
Mexiquc. Bull. Soc. Ent. Fr. 82: 99-104. Description of the imago
(Dlplna: Tcndipedidae). Ent. News 75: 186. First report from
of Diamesa mexicalla n. sp. and D. reissi n. sp.
Cuha.
Sublette, J.E. 1967b. Type specimens of Chironomidae (Diptera)
Roback, S.S. 1964c. Redescription and new records of in the Cornell University Collection. J. K~nsas Ent. Soc. 40:
Col'lO[llIlYPUS humeraUs (Loew). (Diptera: Tendipedidae;
477-564. Description of insulllnus n. gen.; Type species isfurcatus
Pdllpiinac). Notul. Nat. 369: 1-4. New records for Cuba and Johannsen. 193R. Redescription of Chiroflomu.s llnOrl\,mus
1\111,1111;\.
Willi...ron and Trihdo.\ /uguhris (Willi,ton), new combination.
~lItHU'k, S,S. 19658. New ~pccic!. anJ records of Coe/oraflYPu..'i
Suhkttc, J.E. and W. W. Wh1h. 1972. New gencra anu 'Ix:cics
IrlHI1 \1cxico and Central America with a key to the new world
of West Indian ChironomiJac (DipteraL Florid.! Ent. 55: 1-17.
~p("l'Il'S .. outh of the United States (Diptcra: Tendipedidae). Ent.
Description of the imagos of 4 new Xestocilironomlls species from
Nc:w, 76: 29-41. Description of the imago of C. to!tecus n. sp.
Jamaica. Panama and Dominica. 1 Stenochirol1on!us ~pccies from
(~k(i(o) and C. o!m('CWi n. sp. (Mexico. Nicaragua). as well as
Dominica. Petaloc/adius sefoslI.s n. gen .. n. sp. and Arlcy!ocladius
fl'(ords for described species from Panama and Mexico.
relicinus n. gen .. n. sp. from Jamaica. Xcstociliroflomus proposed
Roha<:k, S.S. 1971a. The subfamily Tanypodinae in North as new name for iflSU!WIU5 Sublette. 1967.
Ameri(a (Diptera: Chironomidae). Monogr. Acad. Nat. Sci.
Townes, H.K. 1945. The nearctic species of Tendipcdini (Diptera,
Philadelphia 17:1-410.
Tendipedidae (= Chironomidac). Am. Mid\. Nat. 34: 1-206.
Rohack, S.S. 1971b. A new record and immature stages of Clunio Reports "Chirollomus" (= Coe!dichirollomll.s) carus n. sp. from
f,'(I\iiienJiJ Oliveira. with additional notes on the adults Venezuela. Panama (Canal Zone). Costa Rica; the species is also
(Chironomidae: Clunioninae). Acad. Nat. Sci. Philadelphia. known from Colombia and the U.S.A. (Tc:<.as. Florida).
Nlllulae Nalurae 444: 1-12. New record in Puerto Rico; and some
Utermohl, H. 1958. Zur Gewassertypenfrage tropischer Seen.
en1logi(al data.

437
Verh. Int. Vcr. Limnol. 13: 236-251. Williston, S. W. 1896. On the Diptera of St. Vincent (West Inlilt."'i).
Vargas. L. 19~6.
Jfacropelopia roblesi (Tendipedidae. Diptera) n. Trans. R. Ent. Soc. London 3: 253-276. Description of the ima!-!lls
sp. neotropical procedente de Chiapas. Mexico. Revta Inst. of 9 new species.
Saluhr. Enferm. Trop. (Mexico) 7: 79-84. Description of a Wirth. W. W. 1969. New species and records of Gal~pa~os
prohahly hlood-sucking imago. Diptera. Proc. California Acad. Sci. 36: 571-594. Report of
Vargas, L. 1952. Tendipes (Limnochironomus) califomicus y Thalas'wmyia pi/ipes Edw. from the coast of Baja California.
Tendipes (Limnochironomus) Jigueroai n. sp. (Diptera, Mexico.
Tendipedidae). Revta Soc. Mex. Hist. Nat. 13: 47-51. Ocurrence Wirth, W. W. 1979. Siolimyia amazonica Fittkau. an aquatic
in Mexico. midge new to Florida with nuisance potential. Florida Ent. 62:
Vockeroth, J.R. 1949. Taxonomic studies in the Tendipedidae of 134-135.
Mexico (Diptcra. Nematocera). Masters thesis. Univ. Woke, P.A. 1947. Arthropods of sanitary importance in the
Saskatchewan. 64 p. Republic of Nicaragua. Central America. Amer. J. Trop. Med. 27:
357. Brief mention of three forms identified only to genus.

CERATOPOGONIDAE
Willis W. Wirth

Introduction
The "biting midges" may be found in large numbers in nearly any genera of Neotropical biting midges. Fox (1948-1955) in Puerto
aquatic or semiaquatic habitat in all regions of the world. Because Rico. Floch and Abonnenc (1950) in French Guiana and
of their small size (usually 1-4 mm) they are little collected and Guadeloupe. Vargas (1944-1969) in Mexico. and Wirth and
poorly known compared with mosquitoes or tabanids. Adult habits Blanton (1956-1978) in the U.S.A. published series of revisions of
are diverse but most species are adapted to some type of the Neotropical species of bloodsucking Culicoit/cs. Saunders
bloodsucking. Species of Culicoides. Lasiohelea. and (1956-1964). Soria and co-workers (1975-1976). and Winder and
Leptoconops suck vertebrate blood: some are notorious pests. Silva (1972) have published on the classification and biology of
especially in beach or mountain resort areas. Some Culicoides are the cacao-pollinating midges. Important studies on the biology and
known vectors of diseases. including onchocerciasis of horses and control of Culicoides were made in Brazil by Forattini et al.
cattlc. bluetongue of sheep and cattle. horse-sickness. several (1956-1960) and in Jamaica by Kettle. Linley and Davies
human filariases. and liaemoprott'lH and Leucocyto:oon diseases (1963-1973). Very little work has been done on the biology of
of hirds. In Latin America the pestiferous species are known by the other aquatic ceratopogonids in the region, although there are
common name of 'polvorinos' or 'jejenes.· and in the English scattered papers by Breeland (1960)' Davies and Giglioli (I (79).
speaking West Indies they arc usually called "sand flies." All the Williams (\957. 1964). and Woke (1954). Apparently a large
Ceratopogoninae. with the exception of the tribe Culicoidini. are proportion of the Neotropical species have used aquatic situations
predaceous on soft-bodied insects, especially midges of the family in water-holding plants. tree holes. and similar aerial habitats
Chironomidae. Some Atrichopogoll and Forcipomyia species are (Wirth et al. 1968), possibly an adaptation to escape the severe
ectoparasitic on larger insects. Many genera (e.g. Dasyhelea) are predation and other problems in ground-water biotopes. Wirth and
exclusively flower-visiting and feed on nectar, whereas many Blanton (1959) gave good summaries of the larval habitats and
others supplement their diet or have non-haematophagous species. known adult biologics of the Culicoides of Panama. and in 1974
Species of Forcipomyia and DasyiJelea and small. hairy species of gave a similar treatment for the West Indies species; Aitken et al.
some other ~enera arc important pollinators of cacao (Theobroma (1975) similarly treated the bloodsucking ceratopogonids of
cacao L.) and other tropical crops. Trinidad.
The immature Sla!!CS of Ceratopogonidac may be ferrestrial under Only a small proportion {If thc species of the South and Central
bark or on wet or damp wood (Forcipomyia); semiaquatic on wet American biting midges have been described (Wirth 1474.
alga-covered soil. ""'ood or rocks (A triciJopogon , Dasyhelea. etc.) catalogue). and it is too early to make many generalizations about
or in wet. decomposing plant material such as cactus stems. the fauna and its biogeography. Genera with large numbers of
banana stalks. or leaf compost (Culicoides, etc.); or aquatic in aquatic species are Atrichopogon, Das.vhelea, Culicoides,
mud or wet sand on lake. pond, or stream margins, salt marshes. Monohelea, Stilobe;;;;ia, and Palpom.via. The aql:atic genus
tree holes. or water-holding plants. Most genera have aquatic Be;;zia, very common and widespread in the northern hemisphere.
species which are usually found at or near the water margin or is 'poorly represented in tropical South and Central America. It
water surface where they are associated with the interface of solid seems to have been largely replaced by two very prominent.
substrate. water, and air. related genera. Paryphoconus. which is endemic to tropical South
and Central America. and Stenoxenus, which is pantropical.
The foremost pioneer in the study of Neotropical Ceratopogonidae
was Adolpho Lutz (1912-1914) working at the Instituto Oswaldo Wirth and Blanton (1959) summarized the distribution of the
Cruz in Rio de Janeiro. Brazil. J.W.S. Macfie (1937-1953). for Panama Culicoides according to life zones; they found certain
many years the foremost international specialist in this family. spcics or groups of species were characteristic of each of the
described many Neotropical species. From 1944 to 1961 John humid tropical. arid tropical. subtropical (montane rain forest).
Lane of the Univcrsidade de Sao Paulo, Brazil. revised many and temperate (temperate forest) life zones. Species or groups with

438