Bionimics of black flies (Diptera: Simuliidae) in Costa Rica I Species biting man, with an epidemiological summary for the Western Hemisphere

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(1)Rev. BioI. Trop., 22 ( 1 ) : 187·200, 1 974. Bionomics of black fHes (Diptera: Sirnul'iidae) in C osta Rica. 1. Species biting man, with an epidemiological summary for the Western Hemisphere* by. Mario Vargas V., * * *. B.V. Travis,u. and J . Clyde Swartzwelder* * * * ( Received for pub!ication March 18, 1974) ABSTRACT : In the Western Hemisphere there are listed 61 species of black flies which are recorded as biting rnan in the Neotropical Region and 37 in the Ne:lrctic Region. Five neotropical species, Simulium quadrivillatum, S. metallicum, S. callidum, S. baematopotum and S. paynei, were collected while biting rnln in Costa Rica. The greatest biting rates in Costa Rica were at altitudes of O to 1 500 m.. As part of a program to study the bionomics of biting Diptera In Costa Rica, observations were initiated in JuIy 1968 on the bIack flies. For the first year of the project, observations and collections of bIack flies were made at monthIy intervaIs in 53 streams located in the Central Valley. During the second year, observations were continued at bimonthIy intervals in 28 of these S3 streams and in 47 additional streams located mostly outside the Central Val· ley. It was hoped that the biometrical data assembled from these studies would be helpfuI in a better understanding of the biology and distribution of Louisiana State University International Center for Medical Rese:trch and Training ( LSU·ICMRT) , San ]oEé, Costa Rica. This investigatíon was supported in part by the United States Army Medical Research and Development Cornrnand, Con· tract DADA·17·68 C·8023 . The findings in this report are not to be construed as an official Department of the Arrny position unless so designated by official documents. �' " Louisiana State University, School of Medicine New Orlea!)s, La. Visiting Pro· fessor, on leave from Corne!! University, Ithaca, N.Y. *'lo -' Universidad de Costa Rica, Facultad de Microbiología, Departamento de Parasi. tología, San José, Costa Rica, CA. " " * * Louisiana State University, School of Medicine, New Orleans, La. *. 187.

(2) 188. REVISTA DE BlOLOGIA TROPICAL. species of black flies and that such data would be useful in past and future epidemiological interpretations of disease organisms transmitted by these flies (TRAVIS and VARGAS,. 40;. VARGAS and TRAVI S. ,. 46) .. Because of the general strong reaction to bites and to the transmission. of disease organisms by black flies it seemed appropriate to start assembling lists of black fly species recorded in the Western Hemisphere as biting mano Also it seemed appropriate to make sorne brief cornments on the disease orga nisms transmitted to man by these fIies. In the Western Hemisphere two lists of species are available, VULCANO (47) lists 2 5 3 species in the Neotropical Region, and STONE ( 3 5 ) 1 1 1 species and one subspecies in the Nearctic Regíon ( north of Mexico ) . With the two lists of species as guide lines for valid names, prelimi nary lists were prepared from the literature of species that were recorded as biting mano try.. Each record has one documentation for the species in each coun. In most cases the documentation is not necessarily the original.. nomenc!ature. at the. subgeneric level. is. still. a. matter of. debate,. As the ooly the. generic nap1es are used.. The. species are arranged alphabetically, with the generic names sepa. rated by a comma from the trivial name.. The names of the describers of the. various species are not added in this report as they are induded in the two published Iists.. SPECIES RECORDED BITING MAN NEOTROPlCAL REGION The records for the neotropics indude those published for the Latin American countries, and aIso the records for the biting adults from Oil! Costa Rican studies. In Table 1 are included the. 61 species that are recorded in the litera In tables 2 and 3 are summarized the. ture as biting man in the neotropics.. average numbers of biting adults taken in Costa Rica at various altitudes during. a two-year study. Table 2-A includes the data obtained from collections at 5 3 streams in the Central Valley during the year 1 968-69. The majar objec tives were to concentrate on the larvae, pupae and reared adults. The biting adults collected were. almost exclusively those thal:' were biting the members of the. survey team during the time spent collecting the immatures.. With few exceptions. the total time spent with the abservations of each stream slte was one haur during the period. 9 AM to 1 PM.. There were, with few exceptions, fcm field. workers making the abservations and collections.. Thl,ls, the average numher of. per �bservation 48 man-hours per stream site.. adults per stream site was based on approximately four man-hours and 12 monthly observations, or a total of Table 2-B includes. the Central Valley.. data. from collections at 47 stream sites. Each of these streams was. outside. observed bimonthly for one.

(3) TRAVIS: el al. : BIONOMICS OF BLACK FLIES.. 1. 189. TABLE 1. Black fly species recorded in the literature as biting man in the Neotl'opical Region Species. amazonicum,. Simulium. annulatum, Simulium. Country. References. Venezuela. BRlCEÑO IRAGORRY, VARGAS & DIAZ NAJERA ( 4 ). Argentina. WYGODZINSKY &. Argentina Antilles, Mexico Cuba Me,xico México, Guatemala Colombia Costa Rica. anthracinum, Aust1'osimulium antillarttm, Simulium avidum, Simttlium callidttm, Simttlittm ( =S. mooseri). COSCARON ( 48 ) VARGAS ( 4 2 ) VARGAS (42 ) DIAZ NAJERA ( 1 1 ) HOFFMANN ( 2 0 ). VARGAS ( 4 2 ). GUTTMAN ( 18 ) TRAVIS, VARGAS & SWARTZWELDER*. clarki, Si'lnulium deagostinii, Simttlittlrl delpontei, Simttlittm dilzelli, Simulillm downsi, SimlllimiZ. dureti, Simulium escomeli, Simlllillm exiguum, Simlllillm. flavopllbescens, Simulium ganalesense, Simulium gonzalesi, Simulium gonzalezherrejoni, Simulium guianense, Simulium haematopotum, Simulium. ignescens, Simulium inaequale, Simulium inCI'ustatum, Simulium infuscatllJ1z, Simulium jacumbae, Simulium jerezensis, Simulium juju)'ense, Simulium lahillei, Simulium. •. Panamá Argentina Argentina Argentina Mexico. VARGAS ( 4 2 ) COSCARON ( 6 ) VARGAS ( 42 ) VARGAS ( 42 ) VARGAS & DIAZ NAJERA ( 4 3 ). Guatemala Chile Ecuador Ecuador Colombia Guatemala Brazil Mexico Mexico Mexico British Guiana Mexico Costa Rica. DALMAT ( 7 ). Guatemala Panamá Ecuador Argentina Brazil Brazil 'Guatemala, Mexico Mexico Argentina, Ecuador Argentina. CO SCARON (6). LEON & WYGODZINSKY ( 2 3 ). LEON & WYGODZINSKY ( 2 3 ) BARRETO ( 2 ) DALMAT ( 7 ) VARGAS ( 42 ) VARGAS & DIAZ NAJERA ( 4 3 ) VARGAS & DIAZ NAJERA ( 44) DIAz NAJERA ( 1 3 ). VARGAS (42) VARGAS ( 4 2 ) TRAVIS, VARGAS & SWARTZWELDER* DALMAT (7) FAIRCHILD ( 14 ) LEON & WYGODZINSKY ( 2 3 ) VARGAS ( 4 2 ) VARGAS ( 4 2 ) VARGAS ( 4 ? ). COSCARON ( 6 ) DIAZ NAJERA ( 1 3 ) LEON & WYGODZINSKY BARRETO ( 2 ). (23).

(4) REVISTA DE BIOLOGIA TROPICAL. 190. TABLE 1. ( Cont , ). Elack jly species recorded i n the litl?1'ature. biting m(/n. as. in the Neotl"Opical Region. Species. Country. Brazil Venezuela lugubre, Simulium ( = S, guianense) Venezuela Venezuela lutzianum, Simulium Mexico mediovittatum, Simu/ium lane-portoi, Simulium limbatum, Simulium. meridiona/e, Símu/íum. Mexico. metallicum, Simu/ium. Mexico, Guatemala Costa Rica, Panama Venezuela, Trinidad Colombia BraziJ, Paraguay Costa Rica Chile Mexico, Guatemala Panama, Venezuela Chile Brazil, Paraguay Guatemala Costa Rica Argentina, Brazil, Paraguay, Venezuela Costa Rica. minuseu/um, Simu/ium moorei, Simftlium ochrarellm, Simut;um. opalínijrons, Símll/illm O1"bíta/e, Símlllíum pacheco/unai, Cnephia panamense, Símu/íum. paragllayense, Símll/ium. paynei, Simulillm. References VARGAS ( 4 2 ) VARGAS ( 4 2 ) VARGAS ( 42 ) VARGAS ( 42 ) VARGAS. &.. DIAZ NAJERA ( 45 ). VARGAS (42 ). VARGAS ( 42 ) GUTTMAN ( 1 8 ) VARGAS ( 42 ) VARGAS ( 4 2 ). VARGAS (42 ) COSCARON ( 6 ). COSCARON ( 6 ). DALMAT ( 7 ). ZELEDON & VIETO (4 9 ). COSCARON ( 6 ) TRAVIS, VARGAS & SWARTZWELDER*. Brazil, Argentina Paraguay Brazil Mexico. perti¡lax, Simulium pruinosum, Simlllium. ,. pseudocallidum, Simulium pseudohaematopotum, Simulium. ( = S haematopotum) pulverulentum, Simulillm quadrivittatllm, Simlllium. r¡veti, Simulium. •. Mexico Venezuela. ". VARGAS ( 42 ) VARGAS ( 4 2 ) DIAZ NAJERA ( 1 2 ) HOFFMAN ( 20 ) BRICEÑO IRAGORRY ( 3 ) VARGAS ( 4 2 ). Cuba, Costa Rica Jamaica, Panama Venezuela, Puerto Rico. VARGAS ( 4 2 ). Ecuador. VARGAS ( 4 2 ).

(5) TRAVIS: et al. : BIONOMICS. TABLE 1. OF BLACK FLlES.. Species. Country. rub1'ithorax, Simulium. Brazil, Argentina Venezuela, British Guiana Colombia, British Guiana, Lesser Antilles Colombia Brazil Brazil Chile Mexico Guatemala Venezuela, Canada, U. S., Mexico. semellatum, Simulium seutistriatum, Simulium simplieicolor, Simulium tenuipes, Simulium triv t um, Simulium vera !r� zanum, Simulium l!ersieolor, Simulium vittatum, S'imulium. � J). 191. ( Cont. ). Blaeh fly speeies ,'eeol'ded in the literature in the Neotropieal Region. sanguineum, Simulium. 1. as. biting. man. Rcferences. VARGAS. (42). SMART ( 32 ) VARGAS (42 ) VARGAS ( 4 2 ) VARGAS ( 4 2 ) VARGAS ( 4 2 ) VARGAS ( 42 ) DALMAT ( 7 ) VARGAS ( 4 2 ) VARGAS. (42). year ( 1969-70) by a three-man-team, or approximately 18 man-hours per year. The data. in Table 2 show that outside the Central Valley the greatest biting rates were at altitudes of 500- 1000 m, with the maximum rates insi de the Central Valley at 500-1 500 m. Part of the low biting rates aboye 1 500 m may be a reflection of fewer streams at the higher altitudes. To obtain some data on the annual variability of black fly production and the biting adults, 28 streams wére selected fram the 53 in the Central Valley for .bimanthly observations far a 12-month periodo The data for the first year 1 968-69 (Table 3-A) were obtained by selecting bimonthly data from the monthly observatians for a 12-month periodo These same 28 streams were also observed on a bimonthly basis for the year 1969-70 (TabIe 3-B ) . The 1968-69 observations (Table 3-A ) of these 28 streams were based on six bimonthly observations of each stream site for approximately 24 man-hours. In 1969-70 (Table 3-B) there were only three observers, thus, there was a total of 18 man-hours of observations of each stream site. At least under the conditions fram which the biting adults were obtained, a very low biting rate on man was actually indicated. Perhaps methods can be devised that will be more conducive to higher biting rates. The numbers oE biting adults are surprisingly similar for the two years of observations (Table 3-A and B. ) . In all of our Costa Rican records of biting adults the predominant species were S. quadrit.:ittatum ( 5 34 specimens) and S. metallicum ( 1 60 spe cimens) . Three other species S. callidum ( 7 specimens) , S. paynei ( 1 spe. cimen) and S. haematopotttm (4 specimens) were taken only rarely. The S. quadrivittatum and the S. metalJicum were both taken mostly at altitudes of.

(6) TABLE 2. >_.. V) N·. B/ack f/y adu/ts ( Simulium) collected while biling man during 1968-69 and 1969-70 at diffewlt a/tiludes in Costa Rica. Number collection sites. Altitude. (m). A.. B.. Total flies. Average number of biting adults per site and species q uadriviItafum. melallicum. callidum. paynei. haematopolum. •. 5 3 sites observed in 1968-69 ( Central V alley ) 500-1000. 17. 32. 0.6. 1.1. 0. 1. O. O. 1 00 1 - 1 500. 23. 236. 7.3. 2.8. 0.2. O. O. 1 50 1-2000. 8. 11. 0.4. 1.0. O. O. O. 2001 -2500. 3. O. 0.3. O. O. O. 2 50 1 - 3000. 2. O. O. O. 0.5. O. 47. sites observed in 1969-70 ( outside the Central Valley) 0-500. 32. 72. 2 .0. 0.1. O. O. 0. 1. 5 0 1 - 1000. 13. 147. 9.9. 1.0. 0.1. O. O. O. O. O. O. O. O. 100 1 - 1 500 1 5 0 1-2000. O. O. O. O. O. O. O. 200 1-2500. O. O. O. O. O. O. O. 2 5 0 1 - 3000. O. O. O. O. O. O. O. ¡;: S. en. ..¡ >. O. tn. tl:1. O t-'. O Cl :; ..¡ i=<1 O ::o!. () > t-'.

(7) TABLE 3 Elack fly adults ( Simulium ) collected while biting man during 1968-69 and 1969-70 al different altitudes in Costa Rica. Number collection sites. Altitude (m). Average number of biting adults per site and species Total flies. quad,.ivittatum. metallicmll. ca/lidulll. paynei. haematopotum. ;;i ::... A. �. 1968-69 observations on 28 stream sites selected from the 5 3 streams in the Central Valley. '\. 5. 12. 1 .4. 0.8. 0.2. O. O. 1001-1500. 15. 181. 8.7. 3.3. 0.1. O. O. 1501-2000. 6. 9. 0.5. 1.0. O. O. O. 2001"2500. O. O. O. O. (). O. O. i 501-3000. 2. O. O. O. 0.5. O. 500-1000 •. B.. �.. ". :tll. 1969-70 obseryations on the 28 selected stream sites in the Central Valley. �00-1000. 5. 20. 2.2. 1.8. O. O. O. 1001-1 500. 15. 179. 10.0. 1 .8. 0.1. O. 00. 1 501-2000. 6. 8. O. 1.3. O. O. O. 2001-2500. O. O. O. O. O. O. O. 250 1-30QO. 2. O. O. O. O. O. O. (3. Z O � R en O. >,j. tll t-< > n :><: >T1 t: tT1 !'". >� \O \j.).

(8) ÍUlVISTA. DE BIOLOGIA TROPlCÁi. .. 0-1 500 m with the greatest numbers occurring at altitudes of 1 000 to 1 500 m. The seven specimens of S. callidum were taken principally at altitudes oE 500- 1000 m. The single specimen oE S. paynei was collected at 2630 m and the four S. haematopotum at 60 m. NEARCTIC REGION. In Table 4 are 1isted 37 nearctic speCles which are recorded as biting or annoying man north of Mexico. INCRIMINAT:ED VECTORS. NEOTROPICAL REGION. Onchocerciasis : In the neotropics onchocerciasis was first diagnosed by ( 30 ) in the Pacific coast of Guatemala. That same year BRUMPT ( 5 ) described a new species o f Onchocerca, O . caecutiens, i n Latin Arrierica. MAL DONADO (24) gave a historica1 account oE the disease in the neotropics. At the present time active foei are known to be in Mexico, Guatemala, and Ve nezuela; autoehthonous cases are reported from Colombia and Surinam and pos sibly also in Brazil. There still remain many unanswe!€!d problems on the dis tribution and transmission of Onchocerca to mano For instance, of the many species oE black flies that are recorded as biting man, only about haH a dozen species are reasonably weH documented as transmitters of O. volvulus, the on1y disease organism cIearly shown to be transmitted by b1ack flies. The species that have been incriminated as vectors of Onchocerca in the neotropics are : S. ochraceum, S. metallicum and S. callid"m ( DALMAT, 7 ) . There is some laboratory evidellce that S. mangabeirai (VARGAS and DIAZ NAJE RA, 4 5 ) , S. gonzalezi (=S. exiguum ) , S. haematopotum, and S. veracrtlzanum may also be vectors (GIBSON an,d DALMAT, 1 7 ) . It is interesting that the same most important vector species occur from Mexico in the north to Costa Rica and from Costa Rica to the northern portions oE South America. Although these same species oecur in Costa Rica, there is no report of human onchocerciasis in this country. The importance of S. callí d"m must be reevaluated as a result of the separation of S. pse"docallid"m from the 5. catlid"m complex by . DIAZ-NAJERA ( 12 ) . As might b� expected i n the literature, the certified transmitters oE Onchocerca were in the lists of anthropophi1ic species. However, the fact that a given species of b1ack fly is anthropophilic is no proof that the species will be a transmitter of Onchocerca. It might be assumed tha:t some species have mixed host preferences and normally may be zoophilic. . However, in the absence or in low populations of animal hosts these species may be forced to be strongly anthropophilic. It a1so might be assumed that a reasonably high pop ulation of intermediate hosts such as black flies will be needed to maintain the required transmissions that ;lssure survival of the parasites for which they ROBLES. are vectors.. ;... ..

(9) TRAVIS: el al.: BIONOMICS OF BLACK FLlES.. 1. 195. . TABLIr 4 B/ack f/y species feco"ded in the /ítemture as biting or as pes!s of man 111 the Nearctic Region. Species. Country. R�ferences. arcticum, Simulium. u. S. Canada. PETERSON ( 2 6 ) SOMMERMAN, SAlLER & ESSELBAUGH ( 34 ). aureum, Simulium bicorne, Símu/íum eanon;eo/a, Simulium caudatum, ProsimuUum co" bis, Símulittm decorum, Simulium defo/iarti, Simu/íum doveri, Prosimu/ium esselbaughi, Prosimulium exigens, Prosimulium f/aviantennum, Prosimulium fontanum, Prosim-u/ium fu/vum, Prosimulium fllsmm, Prosimulium birtipes, Prosimu/ium* hunteri, Sim1l/ium ¡en ni"gsi, Simu/ium iohannseni, Simu/i1lm lalipes, Simll/i1lm longisty/atum, Simu/imn luggeri, Simtdium magntcm, ProSÍ'mu/ium meridiona/e', Simu/ium mixtum, Prosimu/ium 1llu/tidentatllm, Prosimu/ium 1ll utata, Cnephia ,¡yssa, Simttlium. parnasSZlm, Simulill1ll pecuarum, Cnepbia ¡,ictipes, Simll/ium pugetense, Simu/ium tescOfum, Simulium t"ivitftttum, Simuliu1ll tuberosmn, Simu/ium 1'entlstum, Simu/ium vittatum, Simu/ium *. u. S. Canada U. S. ( Pest ) U. S. Canada Canada U. S. Canada Canada ( Pest) Canada· Canada U. S. ( Pest) U. S. ( Pest) Canltel'a ( Pese) Canada U. S. Canada U. S. Canada U. S. Canada U. S. Canada U. S : U. S. Canada Canada U. S. Canada U. S. Canada U. S. Canada U. S. ( Pest ) U. S. Canada U. S. U. S. U. S. u. S. u. S. U. S.. Canada U. S. Canada Canada U. S.. STONE & ]AMNBACK ( 37 ) PETERSON ( 2 7 ) PETERSON ( 26 ) PE TERSON ( 27 ) SHEWELL ( 3 1 ) HOCKING & RICHARD S ( 19 ) ANDERSON & DEFOLIART ( 1 ) DAVIES & PETERSON ( 8 ) PE'I'ERSON ( 2 6 ) PETERSONI ( 2·8') SOMMERMAN ( 3 3 ) PETERSON ( 2 7 ) PETERSON ( 26 ) DAVIES� PETERSON & WOOD ( 9 ) PETER<SON ( 2 7 ) ANDERSON &' DEFOLIART ( 1 ) DAVIES & SYME ( lO ) STONE & ]A.i\{ NBACK ( 37 ). HqCKING & RICHARDS ( 19 ) DAVIES & PETERSON ( 8 ) PETERSON ( 2 6 ) STONE & SNODDY ( 3 8 ) DAVIES, PETERSON l,{ \'V'OOD ( 9 ). ]OBBINS-POMEROY ( 2 1 ). STONE & ]AMNBACK ( 37 ) DAVIES, PETERSON & WOOD ( 9 ) FREDEEN ( 16 ) STONE & SNODDY ( 3 8 ) FREDEEN ( 16 ). .. ANDERSON & DEFOLIART ( 1 ) DAVIES & PETERSON ( 8 ) STONE & SNODDY ( 3 8 ) DAVIES & PETERSON ( 8 ) STONE & SNODDY ( 3 8 ) STONE & SNODDY ( 38 ) HOCKING � RICHARDS (' 19 )'. ]OHANNSEN (22 ) : SmNE & SN ODDY .( 3 8 ). S'rONE & ]AMNBACK ( 37 ) STONE & BoREHAM PETERSON ( 2 6 ). ( 36'). STONE & SNODDY ( 3 8 ) DAVIES, PETERSON & WOOD ( 9 ) STONE & ]AM NBACK ( 3 7 ) PETERSON & WEST ( 29 ) TWINN ( 4 1 ) STONE .& SNODDY ( 38). Prosimu/ium hinipes is now thought to be limited to tlie Eastera Herilisphere artd represented in the Western Hemisphere by at least three sibling spe'CÍes, P . fontanum, P. fu/vum and P. mixtum..

(10) 196. REVISTA DE BIOLOGIA TROPICAL. Carate : LE ON and WYGODZINSKY ( 2 3 ) pointed out that in the neo tropics as early as 1862 Juan León of México believed that the disease called "mal de pinto" or "carate" was transmitted by "jejenes" (b1ack fIies) . PEÑA CHAVARRIA and SHIPLEY ( 2 5 ) reviewed the disease and presented some epi demiological evidence that black fIies may transmit the disease caUed carate. The primary evidence is that the disease is distinctly more common in the vicinity of streams and particularly in areas of high black fIy biting rates. The method of transmission of the causative organisin Tre¡JOnema carateum, is still unknown. Venezuelan equine encephalitis (VEE) and vesicular stomatitis: Two additional di seas e organisms have been recently isolated from · black fIies in Colombia. These are the viruses that cause VEE, (TRAPIDO, 39) and vesicular stomatitis (H. TRAPIDO, personal communication ) . The role of black flies in transmitting these viruses is yet to be learned. NEARCTIC REGION. Typhotd and péüagra: In the Nearctic Region no disease is known to be transmitted to man by black fIies, However, in the 1iterature of the early 1900's these insects were accused of transmitting ethio10gic agents causing typhoid and pellagra. Some strong evidence for the transmission of pellagra by b1ack flies in the U.S. was presented by FORBES ( 1 5 ) . Of course we now know that this prob1em is caused by a vitamin deficiency. Equine encephalitÍJ (EE) : Black flies (S. johannsení) were experi mentally infected with EE when fed on infected birds in 'Wisconsin. A1so this virus was iso1ated from a pool of S. merídíonale collectcd from a turkey brooder house ( ANDERSON and DEFOLIART, 1 ) . Black fIy adults vary i n their choice of vertebrate hosts; for instan ce, some species feed most1y on birds. These ornithophilic �pecies may be of con siderable importance in maintaining a virus such as EE by passing the virus from bird to bird. Some species seem to select mammals such as horses and cattle, and some are at least reasonably anthropophilic. RESUMEN. De las especies de simúlidos señaladas en el Hemisferio Occidental co mo picadoras del hombre, 6 1 corresponden a la Región Neotrópica y 37 a la Región Neártica, Durante los dos años dédicados a este estudio, cinco especies t:.eotrop i cales picaron al hombre en alguna oportunidad. De acuerdo con un orden de creciente de antropofi1ia tales especies fueron : Simulium quadtú'ittatum ( 534 especímenes ) , S. metafticum ( 1 60 especímenes ) , S, callidtfm (7 especímenes) , S. haematopotum (4 especímenes) y S. paynei ( 1 esp écimen ) . Dado el inte rés epidemiológico se señala las principales especies de simúlidos incriminadas como vectores, especialmente de oncocercosis, en el Hemisferio Occidental. En Costa Rica la mayor frecuencia de picadas por parte de los simú lidos al hombre se encuentra entre los O y 105 1 500 m sobre el nivel del mar..

(11) TRAVIS:. el. al. : BIONOMICS OP BLACK FUES.. 197. ACKNOWLEDGEMENTS. The authors wish to express their thanks to a number of people for their help with the necessary local arrangements. Special thanks are extended to the Staff of the lQuisiana State University International Center for Medica:l Research and Trainjng (lSU-ICMRT) and also to the laboratory and field assistants, Roberto Echeverri (ICMRT) and Guillermo Zúñiga (Universidad de Costa Rica) . Thanks are also extended to Dr. A. Díaz Nájera of México for kipdly . 1oaning specimens, for reviewing this paper, and fa! supplying additional references. The authars would appreciate being advised by the readers of species not included in Tables 1 and 4 that were missed in the literature review. These records can be sent ta B.V. Travis, Department oí Entomology, Cornell University, Ithaca, N.Y. 14850. LITERATURE CITED 1.. ANDERsoN, ]. R., & G. R. DEFOLIART. 196 1 . 2.. Feeding behavior and host preferences of sorne black flies ( Diptera : Simu liiclae) in Wisconsin. Ann. Elztomol. Soco Amer., 54: 7 1 6-729.. BARRETO, P . .. 1969.. The Species of black flies found. ID. Colombia ( Diptera: SimuJiidae ) . J. N.Y.. Entamo!. Soc., 77: 3 1- 3 5 .. 3.. BRICEÑO I RACORr'1Y, L.. 1943.. Nota sobre SimúJidos del Valle de Caracas. 1 82-186.. Bol.. lAb. Clin . "Luis Ra-. zetti", Caracas, 3 :. 4.. BlUCEÑO IRAGORRY, L., L. VARGA S, & A. DIAZ NAJERA. 1953. 5.. BRUMPT, EMILE. 1919. 6.. The black flies ·( Diptera, Simuliidae ) of Guatemala and tbe;r role as vectors of onchocerC'Íasis. Smithsollian Misc. Collect., 1 2 5 : 1-42 5 .. DAVIES, D . M., & B. V. P ETERSON. 1956. .. 9.. Nuevos datos relacionados con la importancia sanitaria ce los simúlidos. ( Insecta : Diptera) . Rev. Soco Entamo!. Ar., 3 1 : 7 3-76.. DALMAT, H . T.. 1955.. 8.. Una nouvelJe filaire patbogene parasite de l'homme ( Oncboc8rca caecl/tiens, n. sp.) . . BI/II. Soco Pa/h. Exot., 1 2 : 464-473.. COSCARON, S.. 1 969. 7.. Algunos caracteres morfológicos de Siml/lit/m amtlzonictlm Goeldi, 1905 ( Insecta, Diptera ) . Re !!. Il1st. Stdl/br. Enferm. Tro/J. ( Méx. ) , 1 3 : 243-245.. Observations on tbe mating, feeding, ovarian development and oviposition of � dult black flies ( Simuiiidae: Diptera ) . Can. J. Zool., 34: 6 1 5-65 5 .. DAVI E s, D . M., B . V . PETERSON, & D . M . WOOD 1962. The black flies (Diptera : Simuliidae) of Ontario. 1. Adult identification and .distribution with descriptions of six new species. Proc. Entamo!. Soco 0111., 92 : 70- 154..

(12) REVISTA. 198 10.. DE. mOLOGIA TROPI.CAL. DAVIEs, D. M., & P. D. SYME 1958. Three new Ontario black flies of the Genus Prosimulium ( Diptera : Simuliidae) . n. Ecological observations and experiments. Can. Enlomol., 90 : 744-759.. 11.. DIAz NA.JERA,. 1961.. 12.. 13.. DIAl NAJERA, A. 1965. Acerca de un nuevo simúlido mexicano (Diptera: Símuliidae') . Salubr. Enferm. Trop. (Méx. ) , 2 5 : 2 5 -45 .. Simúlidos de Aguas Calientes y Zacatecas. Descripción de. s.eis esp.ecies .del subgénero Psilopelmia y datos de dimorfismo. Rev. Inves/. Salud Pública, (Méx. ) , 29: 2 1-63.. FAIRCHI LD, G. B.. 1940.. Notes on the Simuliidae of Panamá. Soco Amer., 33: 701-7 19.. (Dipt., N ematocera ) . Ann.. 15.. FORBES, S. A. 1 9 1 2 . O n black flies and buffalo gnats ( Simu/illm ). 1 6.. FREDEEN, F . ]. H.. Entomo/.. as possible carriers of pelJagra in Illinois. 27th. Rept. Sta. Entomol. Ill. Bull ., 1 5 9 : 2 1-52.. 1958.. 17.. Rev: Inst.. DIAZ NAJERA, A.. 1969.. 14.. A. Nota acerca de Simulium (Psilopelm ia ) antillartlm Jennings, 1915, nueva especie para México (Diptera: Simuliidae) . Rev. Iml. 5alubr. Enferm·. Trop. (Méx. ) , 2 1 : 79-9 2 .. Black flies (Diptera: Simuliidae) of the agricultural areas of Manitoba, Saskatchewan, and Albert:!. Pro c. Tenth Int. Congr. Enromol., 3 : 819-823 .. GIBSON, e, & H. DALMAT. 1952.. Three new potential intermediate hosts of human onchocerciasis in Guate mala. A mer. ]. Trop. Med. Hyg., 1 : 848-85 1 .. 18.. GUTTMAN, D. 1972. The biting actmty of black flies: ( Diptera : Simuliidae). in three types of habitats in western Colombia. J. Med. Entomol., 9: 269-276.. 19.. HOCKING, B., & W. R. RICHARDS. 1952.. 20.. Biology and control of Labrador black flies (Diptera : Simuliidae) . Bull . Entomo/. Res., 4 3 : 2 37-257.. HOFFMANN, e. 1930.. C, Los simúlidos de la reglOn onchocercosa de Chiapas ( con descripción de nuevas especies) . An. Insl. Bio!. (Méx. ) , 293-306 . ... 2 1.. 1 9 1 6.. 22.. A. W. Notes on five North American buffalo gnats of t�1e genus Simulium U.S.D.A. B ull ., 329: 1-48.. ]OBBINS-POMEROY,. ]OHANNSEN, O.. 1934.. A. Aquatic Diptera. 1. Nematocera, exclusive of Chironomidae and Ceratopo gonidae. Co rnell Uni v. Agr. Exp. Sta. Memoir, 164·: 1-7 1 ..

(13) TRAVIS: el al.: BlONOMlcS OF BUCK FLlES. 23.. LEON,. l'9H. 24.. L. A., & P. WYGODZINSKY Los simúlidos del Ecuador (Diptera) .. He/mintiasis del Hombre en América.. lona. 25.. 199. Bol. Inf. Ciento Nac, 6:. 269-288.. F.. MALDONADO, J.. 1965.. I. Editorial Científico-Médica, Barce·. 529 pp.. A., & P. G , SHIPLEY Contribución al estudio de los carates de América Tropical. Rev. Med. Latino. PEÑA CHAVARRIA,. 1925.. Amer.,. 10:. 3-76.. B. V. Observations on mating, feeding and oviposlt1on of sorne Utah species of black flies (Diptera: Simuliidae) . Can. Entomo/., 9 1 : 147- 1 5 5 .. 26. ' PETERSON, 1959.. 27.. 1959.. 28.. B. V. Notes on the biology oE sorne species oE Utah blackflies ( Diptera: Simu1iidae) . Mosq. News, 1 9 : 86-90.. PETERSON,. B. V. The Prosimll/illm of Canada and Alaska. PETERSON,. 1970.. Entomo/. Sor. Can., 29.. Enfermedad de Rob/es. Academia de Ciencias Médicas, Físicas y Naturales, 1963. P. 58·82.. SHEWELL, G. E.. 1959.. New Canadian black flies ( Diptera: Simuliidae). Simuliidae (Dipt.) from British Guiana and the Lesser Antilles. Roy. Entomo/. Sor. London,. SOMMERMAN, K.. 1964.. 34.. Can. Enlomo/., 9 1 : 686·697.. S�ART, .J.. 1940.. 33.. Mem.. D. G., & A. S. WEST Control of adult black flies ( Diptera : Simuliidae) in the forests of Eastero Canada by aircraft spraying. Can. Enlomo/., 9 2 : 7 14·719.. Guatemala ( ed.),. 32.. Simuliidae) .. ROBLES, R.. 1919.. 31.. ( Diptera :. 1 · 2 1 6.. PETERSON,. 1960.. 30.. 69:. 90:. Trans.. 1·10.. M.. Prosimlllillm eJse/ballghi n. sp., the Alaska P. hirlipes 2 . liidae) . Pror. Entomo/. Sor. ' Wash., 66: 141· 145.. SOMMERMAN, K.. 1955.. M., R. I. SAILER, & C. O. ESSELBAUGH Biology of Alaskan black flies (Simuliidae, Díptera) .. ( Diptera: Simu. Ero/. Monogr., 2 5 :. 345-385. 35.. STONE,. 1965.. 36.. M. M. BOREHAM A new specíes of Simll/illm from the southwestern United States (Diptera: Simuliidae) . ]. Med. Entomo/., 2 : 1 64-170 .. STONE, A., &. 1965.. 37.. A. Family Simuliidae, p . 1 8 1 - 1 89. In A catalog of the Díptera 01 America norlh 01 Mexico. U.S.D.A. Handbook 276.. STONE, 1955.. A., & H.. A.. ]AMNBACK. The b/adz flies of New Bull ., N9 349. 144 pp.. York Sta/e (Diptera: Simllliidae) .. N.Y. State Mus..

(14) REVISTA DE BIOJ.OGIA TROPICAL. 200. 3 8.. STONE,. 1 969.. A., & E. 1. SNODDY The black flies of Alabama Expt. Bull.,. 39.. TRAPill O ,. 1 97 1 .. ( Diptera: Simuliidae) .. Univ.. Aubl¡r11. Agr.. 380 : 1 -9 3 .. H.. Venezuelan eilcephalitis. Proc. encephalitis.. o f the. wOl'kshop .sYlllp osittm. 011. Venezuelall. PAHO Scientific pubJication NQ 243. Washington, D.C. pp.. 1 63-164. 40.. B. V., & M. VARGAS V. Bionomics of black flies. TRAVIS,. 1972.. Costa Rica.. 10. J.. PI·OC. 57th Ami. Meetillg N.. 41.. An ecological. C. R. black flies of eastero Canada Seco D., 1 4 : 97-150.. consideration.. 1 1 1- 1 1 2. TwINN,. 1936.. 42.. II. Mosq. Extel·m. Assoc. :. The. ( SimuIiidae, Diptera ) .. VARGAS, 1. 1945.. SimúlidoJ del Nuevo Mundo. (Méx. ) . 241. p.. Monografía. N° 1. Can.. f.. Res.. del Inst. Salubr. Enf. Trop.. 43.. VARGAS, L., & A. DIAZ NAJERA 1 95 1 . Nota sobre los simúlidos de México y su distribución geográfica (Diptera : Simuliidae) . Re1). Inst. Salubl'. Enf. Trop. ( Méx. ) , 1 2 : 89- 1 00 .. 44.. VARGAS, L., & A. DIAZ NAJ ERA 1953.. Símulium (Notolepl'ia ) f,ollzalesi Enf. Trop. (Méx. ) , 1 3 :. 45.. L., & A. DIAZ NAJ E RA Simúlidos Mexicanos. Rev.. Inst. Salubl'. En!. TI·op.. (.Méx. ) ,. 1 7 : 143-399.. VARGAS V., M., &. 1973.. B. V. TRAVIS Bionomía de los simúlidos (Diptera: Simuliidae) en Costa Rica. IV. Lo calización y descripción de los lugares de recolección. Rev. Biol. Trop., 21:. 47.. Rev. Itm. Salubr.. VARGAS,. 1957. 46.. n. sp . ( Insecta, Diptera ) .. 2 35-239.. 143-175 .. A. Family Simuliidae, p . 1-44. I n. VULCANO, M .. 1967.. south of the United Sta/es.. A catalogue o f the Díptera o f the Americas. 16.. Dept.. Zoo1., Secreto Agrie. Sao Paulo,. Brazi1. 48.. P., & S. COSCARON A review of Simulium ( Pte1'1lctspatha) EnderIein. WYGODZINSKY,. 1967.. Bull. Amer. Mus. Nat. Hist.,. 49.. Diptera) .. R., & P . 1. VIETO Contribución al estudio de los Simúlidos de Costa Rica ( Diptera: Nema tocera ) . I. Sobre el Simulium panamense Fairchild, 1940. Rev. Biol. Trop.,. ZELEDON,. 1957.. ( Simuliidae,. 1 36 : 49- 1 16.. 5 : 19-33..

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