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Si e) pe oO =z | as * _, SMITHSONIAN_INSTITUTION a +, a “oy “ee 6 aie Zz i | NVINOSHLINS S3tYVUaIT wy ne z = YY; = + g fp, 5 6p) vw, WA oO y Seys Fi Hm 2 ; be | a = ; 2 | fa? SMITHSONIAN INSTITUTION NOILN SAiuvustT LIBRARIES Ww eee z ~ & mc Lo i a om.” it. pie © wey 2 i = wo 2) | eee z : pm pe 8 | — os a Zz INSTITUTION NOILNLILS LIBRARI Saacatt \ Fk 2th i + ce Sa ay Det hae Na Contributions of the American Entomological Institute Volume 4, Number 1, 1968 “"v" MOSQUITO STUDIES (Diptera, Culicidae) X. Dixinae originally described from North America. By By T. Michael Peters XI. Mosquitoes originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay. By John N- Belkin, Robert X. Schick, and Sandra J. Heinemann CONTRIBUTIONS of the AMERICAN ENTOMOLOGICAL INSTITUTE The Contributions are for larger papers on insects. Each paper is a separate number, with separate pagination and index. Separate numbers aggregating about 500 pages constitute a volume. Issues appear irregularly, as suitable manuscripts are available. Copies are sold separately or in subscriptions to complete volumes. Complete volumes are $12.00. The price of separate numbers varies. Subscribers are billed for each volume with its beginning number, and receive the parts as issued. Orders for separate numbers that total less than $8.00 must be accompanied by payment. Address orders or correspondence to the American Entomological Institute, 5950 Warren Road, Ann Arbor, Michigan 48105, U. S. A. No. No. No. bide 2a. 3. 2 44 Parts of Volume 1, with prices Dasch, Clement E., 1964. The Neotropic Diplazontinae (Hymenoptera, Ichneumonidae). 75 pages, 69 figures. Price: $2.25, postpaid Mosquito Studies (Diptera, Culicidae), 1965. I. Belkin, John N., etal. A project for a systematic study of the mosquitoes of Middle America. II. Belkin, John N., et al. Methods for the collection, rearing and preservation of mosquitoes. 78 pages, 4figures. Price: $2.25, postpaid Same as no. 2, but in Spanish. Price: $2.25, postpaid Matthews, Robert W., 1965. The biology of Heriades carinata Cresson. (Hymenoptera, Megachilidae) 33 pages, 23 figures. Price: $1.00, postpaid Mosquito studies (Diptera, Culicidae), 1965. Ill. Ramalingam, Shivaji, and John N. Belkin. Two new Aedes from Tonga and Samoa. IV. Belkin, John N. The mosquitoes of the Robinson-Peabody Museum of Salem expedi- tion to the southwest Pacific, 1956. 34 pages, 3figures. Price: $1.00, postpaid Mosquito studies (Diptera, Culicidae), 1965. V. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes originally described from Middle America. 95 pages. Price: $2.75, postpaid Mosquito studies (Diptera, Culicidae), 1966. VI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes originally described from North America. 39 pages. Price: $1.00, postpaid DeLong, Dwight M. and Paul H. Freytag, 1967. Studies of the world Gyponinae (Homopotera, Cicadellidae). A synopsis of the genus Ponana. 86 pages, 257 figures. Price: $2.50, postpaid Contributions to the mosquito fauna of southeast Asia. I. Delfinado, Mercedes D. The genus Aedes, subgenus Neomacleaya Theobald in Thailand. 55 pages, 20 figures. Price: $1.75, postpaid Parts of Volume 2, with prices Contributions to the mosquito fauna of southeast Asia, 1967. Il. Bram, Ralph A. The genus Culex in Thailand (Diptera: Culicidae). 296 pages, 104 figures. Price: $9.00, postpaid Schuh, Toby, 1967. The shore bugs (Hemiptera: Saldidae) of the Great Lakes region. 35 pages, 30 figures. Price: $1.25, postpaid Snider, Richard J., 1967. The chaetotaxy of North American Lepidocyrtus s. str. (Collembola: Entomobryidae). 28 pages, 57 figures. Price: $1.00, postpaid — Contributions to the mosquito fauna of southeast Asia. : Ill. Delfinado, Mercedes D., 1968. The genus Aedes, subgenus Neomacleaya Theobald of southeast Asia. 76 pages, 26figures. Price: $2.25, postpaid Contributions to the mosquito fauna of southeast Asia. IV. Knight, Kenneth L., 1968. Species of the subgroup Chrysolineatus of group D, genus Aedes, subgenus Finlaya Theobald. 45 pages, 12 figures. Price: $1.50, postpaid Parts of Volume 3, with prices Mosquito studies (Diptera, Culicidae), 1968. VII. Belkin, John N. The Culicidae of New Zealand. 182 pages, 30 figures. Price: $5.50, postpaid Mosquito studies (Diptera, Culicidae), 1968. VIll. Zavortink, Thomas J. A prodrome of the genus Orthopodomyia. 221 pages, 41 figures. Price: $6.75, postpaid Fleming, Richard C., 1968. Head musculature of sphinx moths (Lepidoptera: Sphingidae). 32 pages, 37 figures. Price: $1.00, postpaid Mosquito studies (Diptera, Culicidae), 1968. IX. Belkin, John N. The type specimens of New World mosquitoes in European museums. 69 pages. Price: $2.00, postpaid Parts of Volume 4, with prices Mosquito studies (Diptera, Culicidae), 1968. X. Peters, T. Michael. Dixinae originally described from North America. 7 pages. XI. Belkin, John N., Robert X. Schick, and Sandra J. Heinemann. Mosquitoes originally described from Argentina, Bolivia, Chile, Paraguay, Peru, and Uruguay. 29 pages. Price of X and XI: $1.25, postpaid MOSQUITO STUDIES (Diptera, Culicidae) X. DIXINAE ORIGINALLY DESCRIBED FROM NORTH AMERICA! By T. Michael Peters“ Belkin, Schick and Heinemann (1966) presented all the available data on the source of the type material for all the nominal species of Culicinae and Chaoborinae originally described from North America north of Mexico. In the present paper the treatment of the family Culicidae in the broad Sense is completed for North America by similar information for the Dixinae. The data are presented in the same manner. Included in the list are species described by Peters in Peters and Cook (1966), whose taxonomic treatment is followed here. LIST OF SPECIES Dixinae The habitats of the immature stages of the Dixinae have been recorded for only 16 species in North America. This information is given below under BIO- NOMICS for each of these species. For all the other species the following in- formation is presented here as a guide to locating breeding sites. In general the larvae and pupae are found among vegetation or debris along the margins of various bodies of ground water. They are particularly abundant on the margins of very small streams, springs or seepages but occur also in pools in larger, swifter streams aS well as in permanent, semipermanent and temporary stag- nant pools, ponds, lakes and even in bogs and Swamps. Occasionally they may be present in artificial containers. 256. Dixa arge Dyar & Shannon, 1924b:199-200. TYPE: Holotype o, Longmire Springs (Mt. Rainier National Park), Washington, 14 June 1917, H.G. Dyar (USNM, 27454). BIONOMICS: [Larvae probably among emergent vegetation in clear, eurythermous, moderately fast, acidic to neutral, shallow streams or small rivers. | Pecntibation from project ''Mosquitoes of Middle America" supported by Public Health Service Research Grant AI-04379 and U.S. Army Medical Re- search and Development Command Research Contract DA-49-193-MD-2478. Ee et of Entomology, University of Massachusetts, Amherst, Mass. | 01002. 2 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 257. Dixa blax Dyar & Shannon, 1924b:199. TYPE: Holotype “, Bright Angel (Coconino Co.), Arizona, 10 May 1903, H.S. Barber (USNM, 27453). 258. Dixa brevis Garrett, 1924a:6. TYPE: Holotype “, McLean Creek, about 2 mi from mouth, Okanagan Falls, British Columbia, 23 Sept, C. Garrett (CNC). BIONOMICS: Males occur in swarms flying low along the banks about a foot off the water. [Larvae probably in shallow acidic seepage pools or hoof- prints. | 259. Dixa distincta Garrett, 1925a:12. TYPE: Holotype & (130), St. Joseph Creek, Cranbrook, British Columbia, 16 Oct, C. Garrett (CNC). 260. Dixa fluvica Peters, 1966:242-243. TYPE: Holotype “, Valley Creek (Washington Co.), Minnesota, 3 July 1962, T.M. Peters (USNM, 67268). BIO- NOMICS: Adults swept from vegetation near stream. Larvae among duckweed and watercress in quiet backwater just below entrance of spring into clear, fast, rocky stream. 261. Dixa fraterna Garrett, 1924a:6. TYPE: Holotype “, Bull River, British Columbia, 10 Oct 1923, C. Garrett (CNC). 262. Dixa fusca Loew, 1863a:4. TYPE: Lectotype 2, New York, probab- ly collected by Osten Sacken (MCZ 10391, by PRESENT SELECTION; MCZ 10391 [2] here designated as paratebtatype). 263. Dixa mystica Dyar & Shannon, 1924b:197-198 [=fusca]. TYPE: Lec- totype &, Cabin John (Montgomery Co. ), Maryland, 22 Feb 1915, R.C. Shannon (USNM, 27451, by PRESENT SELECTION; & adult, Difficult Run (Fairfax Co.), Virginia, 3 Oct 1915, W.R. McAtee, USNM, 27451 is here designated paralec- totype). 264. Dixa hegemonica Dyar & Shannon, 1924b:194-195. TYPE: Holotype co, Eureka (Humboldt Co.), California, 25 May 1903, H.S. Barber (USNM, 27446). 265. Dixa inextricata Dyar & Shannon, 1924b:198-199. TYPE: Lectotype o, (slide 2008), Dead Run (Fairfax Co.), Virginia, 25 Mar 1914, R.C. Shannon (USNM, 27452; selection of Peters and Cook, 1966:245). 266. Dixa johannseni Garrett, 1924a:7. TYPE: Holotype “, Michel, Brit- ish Columbia, 21 Sept, C. Garrett (CNC). BIONOMICS: [Adults probably swarm above streams. | 267. Dixa lobata Garrett, 1924a:6. TYPE: Holotype “, Cranbrook, Brit- ish Columbia, 30 June, C. Garrett (CNC). 268. Dixa melanderi Peters, 1966:246. TYPE: Holotype , Puget, Wash- ington, 4 July 1924, A.L. Melander (USNM, 67269). 269. Dixa modesta Johannsen, 1903b:429-431. TYPE: Lectotype <, Ithaca (Tompkins Co.), New York, Apr 1902, O.A. Johannsen (Cornell 2992, by PRESENT SELECTION). BIONOMICS: Larvae in slow flowing streams. 270. Dixa neohegemonica Peters, 1966:247. TYPE: Holotype “, Mill Valley (Marin Co.), California, 6 May 1936 (CAS). 271. Dixa notata Loew, 1863a:4. TYPE: Lectotype 2, Maryland, probab- ly collected by Osten Sacken (MCZ 10393, by PRESENT SELECTION; MCZ 10393 [2] is here designated paralectotype). 272. Dixa pullogruma Peters, 1966:248-249. TYPE: Holotype 0, Rapid Creek, 2 mi E of Rochford (Pennington Co.), South Dakota, 14 Sept 1961, T.M. Peters et al. (USNM, 67270). BIONOMICS: Larvae ina fast, cold stream. 273. Dixa repanda Peters, 1966:249. TYPE: Holotype “, Goodhue Co., Minnesota, 21 May 1937, D.G. Denning (USNM, 67271). 274. Dixa rhathyme Dyar & Shannon, 1924b:197. TYPE: Holotype ~, Glacier (Whatcom Co.), Washington, 4 June 1917, H.B. Dyar (USNM, 27450). Peters: Topotypic North American Dixinae a 275. Dixa montana Garrett, 1924a:5; garretti Cooper & Rapp, 1944:247, new name [=rhathyme|. TYPE: Holotype , Cranbrook, British Columbia, 30 June, C. Garrett (CNC). 276. Dixa parva Garrett, 1924a:6. [=rhathyme]. TYPE: Bee 7. Wasa, British Columbia, Oct, C. Garrett (CNC). 277. Dixa rudis Garrett, 1924a:6. TYPE: Holotype ¢, Ta Ta eee? Wasa, British Columbia, 27 Sept, C. Garrett (CNC). 278. Dixa similis Johannsen, 1923a:57-58. TYPE: Holotype “, Ithaca (Tompkins Co.), New York, Aug, O.H. Johannsen (Cornell 2993). 279. Dixa terna Loew, 1863a:3. TYPE: Lectotype o, New York, probab- ly collected by Osten Sacken (MCZ 10394, by PRESENT SELECTION: MCZ 10394 [2] is here designated as paralectotype). 280. Dixa venosa Loew, 1872a:50. TYPE: Holotype 2, Belfrage, Texas -(MCZ 10395). 281. Dixa xavia Dyar & Shannon, 1924b:195-196. TYPE: Holotype “, Los Gatos (Santa Clara Co.), California, 16 Feb 1906, J.M. Aldrich (USNM, 27448). 282. Dixella aliciae (Johannsen, 1924b:45-46). TYPE: Holotype &, Clare- mont, across street E from High School (Los Angeles Co.), California, 1 May 1923, Alica M. Westfall (Cornell 2995). BIONOMICS: Reared from larvae collected from surface of pool formed by leak in irrigation pipe. 283. Dixella aurora Peters, 1966:254-255. TYPE: Holotype “, swamp, Ringwood Reservoir (Tompkins Co.), New York, 16-17 June 1963, W.W. Wirth (USNM, 67272). 284. Dixella californica (Johannsen, 1923a:54-55). TYPE: Holotype &, Stanford University (Santa Clara Co.), California, Mar (Cornell 2770). BIO- NOMICS: [Larvae probably in clear, shallow, eurythermous, acidic pools with muddy bottoms and emergent vegetation. | 285. Dixella thones (Dyar & Shannon, 1924b:193-194) [=californica]. TYPE: Lectotype oc’, Longmire Springs [Mt. Rainier Nat. Park], Washington, 2 Aug 1905, J.M. Aldrich (USNM, 27445, slide 2028, by PRESENT SELEC- TION; USNM, 27445, 14 June 1917, H.G. Dyar, is here designated paralecto- type). 286. Dixella clavata (Loew, 1869b:2). TYPE: Lectotype 2, Lenox (Berk- shire Co.), Massachusetts, Osten Sacken (MCZ 10390, by PRESENT SELEC- TION; MCZ 10390 [2] is here designated a paralectotype. ) 287. Dixella cornuta (Johannsen, 1923a:55-56). TYPE: Holotype “, Mc- Lean (Tompkins Co.), New York, Oct (Cornell 2990). BIONOMICS: [Larvae probably in permanent open or temporary woodland ponds, acidic to neutral, less than 6 feet deep, 11-28°C, with muddy or leafy bottom. May also occur in slow to fast, shallow, slightly acidic, 14-22°C stream. | 288. Dixella plexipus (Garrett, 1925b:5) [=cornuta]. TYPE: Holotype & (150), Nelson, British Columbia, June, C. Garrett (CNC). 289. Dixella simplex (Garrett, 1925a:12) [=cornuta]. TYPE: Lectotype 3, Cranbrook, British Columbia, June, C. Garrett (CNC; by PRESENT SE- LECTION). 290. Dixella somnolenta (Dyar & Shannon, 1924b:195) [=cornuta]. TYPE: Holotype &, Viola (Latah Co.), Utah, 21 Aug 1912, J.M. Aldrich (USNM, 27447). 291. Dixella spiralis (Garrett, 1924a:1-2) [=cornuta]. TYPE: Lectotype o, Wasa, British Columbia, 22 Oct 1923, C. Garrett (CNC 5332, by PRESENT SELECTION; o, Wasa, British Columbia, 25 Oct 1923 (CNC 5341); o&, Wasa, British Columbia, 21 Oct 1923; o', Wasa, British Columbia, 23 Oct 1923; 4 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 o, Cranbrook, British Columbia, 2 July 1920 (CNC 2618); 0, Cranbrook, Brit- ish Columbia, 21 Oct 1923 (CNC 5511) are here designated paralectotypes). 292. Dixella deltoura Peters, 1966:257-258. TYPE: Holotype 0, Nenana, Alaska, 13 June 1951, R.I. Sailor (USNM, 67274). 293. Dixella dorsalis (Garrett, 1924a:4). TYPE: Holotype &, Wasa, Brit- ish Columbia, 23 Oct, C. Garrett (CNC). BIONOMICS: [Larvae probably in shallow bogs, seepage pools, temporary woodland pools, or permanent open ponds with a temperature range of 13-239C, muddy or leafy bottoms, and 5.0- 6.2 pH. Larvae probably situated at bases of sphagnum or sedge hummocks. Larvae may also occur in large, slow streams less than 4 ft. deep, with a pH range of 6.2-7.0, clear, and with a muddy or sandy rock bottom. | 294. Dixella dyari (Garrett, 1924a:3). TYPE: Holotype 0, Cranbrook, British Columbia, 25 June, C. Garrett (CNC). 295. Dixella indiana (Dyar, 1925:217-218). TYPE: Holotype o, Fort Ben- jamin Harrison (Marion Co.), Indiana, 23 June 1925, H.K.B. Hufford (USNM, 28419). BIONOMICS: [Larvae probably in clear to slightly turbid seepage pools in Slow to moderately fast, clear to rust-colored streams with a range of tem- perature of 15.5-23.5°9C, pH 5.5-8.0, depth of 10 inches-5 feet, muddy to rocky bottom. Adults probably swarm at edges of stream at twilight. | *296. Dixella marginata (Loew, 1863a:2). TYPE: Holotype 9, District of Columbia, Osten Sacken (MCZ 10392). 297. Dixella neoaliciae (Garrett, 1924a:2-3). TYPE: Holotype “, Boze- man (Gallatin Co.), Montana, 3 Sept, C. Garrett (CNC). BIONOMICS: [Larvae probably in eddies of cold, clear, slightly acidic streams or in warm, clear, Slightly acidic ponds less than 3 feet deep and with silted bottom. Adults prob- ably rest in emergent vegetation during the day. | 298. Dixella nova (Walker, 1848:85). TYPE: o, New York Factory, [Hud- son's Bay at mouth of Hayes River, Manitoba] (BM). BIONOMICS: [Larvae probably in shallow, rust-colored bogs and clear temporary woodland pools or seepage pools with leafy bottoms, a temperature range of 13-17. 5°C and pH of 6.0-6. 8. | 299. Dixella centralis (Loew, 1863a:3) [=nova]. TYPE: Holotype 0, New York, Osten Sacken (MCZ 10389). Terminalia of the type is missing. 300. Dixella nocheles (Dyar & Shannon, 1924b:196-197) [=nova]. TYPE: Holotype o, Biscayne Bay (Dade Co.), Florida, probably A. T. Slossen (USNM, 27449). 301. Dixella occidentalis (Garrett, 1924a:4-5) [=nova]. TYPE: Holotype o, Gold Creek, Cranbrook, British Columbia, 1 May, C. Garrett (CNC). 302. Dixella serrata (Garrett, 1924a:2). TYPE: Holotype o, Oliver, Brit- ish Columbia, 6 May, C. Garrett (CNC). BIONOMICS: [Larvae probably in shallow ponds or very slow-moving streams with silt bottoms, a temperature range of 14.5-28°C and pH of 6.8-7.0. Infrequently larvae may be found in clear, fast, warm, rocky, very acidic streams. | 303. Dixella techana Peters, 1966:263. TYPE: Holotype “, Hassayampa River, Wickenburg (Maricopa Co.), Arizona, 29 June 1953, W.W. Wirth (USNM, 67275). BIONOMICS: [Larvae probably in clear, warm, alkaline, shallow stream with current less than 50 ft/min. and a rocky bottom. | 304. Dixella universitatis (Cockerell, 1926:166). TYPE: Holotype §, Boulder (Boulder Co.), Colorado, 27 Sept 1926, Scott Gale (USNM, 29540). 305. Dixella vespertina Peters, 1966:264. TYPE: Holotype “, McFadden Pond, Corvallis (Benton Co.), Oregon, 17 Sept 1953, V. Roth (Oregon State Univ. ). Peters: Topotypic North American Dixinae 3) *306. Meringodixa chalonensis Nowell, 1951:299. TYPE: Lectotype <, Chalone Creek [Pinnacles Nat. Monument], California, 20 Apr 1950, W.R. Nowell (USNM, by PRESENT SELECTION). BIONOMICS: Larvae in protected, shaded spots among rocks in stream. Nomen Dubium 307. Dixa recens Walker, 1848:85. TYPE: ¢, New York Factory [Hud- son's Bay at mouth of Hayes River, Manitoba]; type lost and the description is inadequate for determination of the taxonomic species involved (Peters and Cook, 1966:251). LIST OF LOCALITIES Canada BRITISH COLUMBIA Bull River: 261. Dixa fraterna. Cranbrook: 259. Dixa distincta; 267. Dixa lobata; 294. Dixella dyari; 275. Dixella garretti; 301. Dixella occidentalis; 289. Dixella simplex; 291. Dixella Spiralis. Michel: 266. Dixa johannseni. Nelson: 288. Dixella plexipus. Okanagan Falls: 258. Dixa brevis. Oliver: 302. Dixella serrata. Wasa: 276. Dixa parva; 277. Dixa rudis (Ta Ta Creek); 293. Dixella dor- salis. MANITOBA New York Factory (Hudson's Bay at mouth of Hayes River): 298. Dixella nova; 307. Dixa recens. United States ALASKA Nenana: 292. Dixella deltoura. ARIZONA Bright Angel (Coconino Co.): 257. Dixa blax. Wickenburg, Hassayampa River (Maricopa Co.): 303. Dixella techana. CALIFORNIA Chalone Creek (San Benito Co.): 306. Meringodixa chalonensis. Claremont (Los Angeles Co.): 282. Dixella aliciae. Eureka (Humboldt Co.): 264. Dixa hegemonica. Los Gatos (Santa Clara Co.): 281. Dixa xavia. Mill Valley (Marin Co.): 270. Dixa neohegemonica. Stanford University (Santa Clara Co.): 284. Dixella californica. 6 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 COLORADO Boulder (Boulder Co.): 304. Dixella universitatis. DISTRICT OF COLUMBIA Washington: 296. Dixella marginata. FLORIDA Biscayne Bay (Dade Co.): 300. Dixella nocheles. IDAHO | Viola (Latah Co.): 290. Dixella somnolenta. INDIANA Fort Benjamin Harrison (Marion Co.): 295. Dixella indiana. MARYLAND Cabin John (Montgomery Co.): 263. Dixa mystica. Locality not specified: 271. Dixa notata. MASSACHUSETTS Lenox (Berkshire Co.): 286. Dixella clavata. MINNESOTA Valley Creek (Washington Co.): 260. Dixa fluvica. Goodhue Co.: 273. Dixa repanda. MONTANA Bozeman (Gallatin Co.): 297. Dixella neoaliciae. NEW YORK Ithaca (Tompkins Co.): 269. Dixa modesta; 278. Dixa similis. McLean (Tompkins Co.): 287. Dixella cornuta. Ringwood Reservoir (Tompkiys Co.): 283. Dixella aurora. Locality not specified: 262. Dixa fusca; 279. Dixa terna; 299. Dixella centralis. OREGON Corvallis, McFadden Pond (Benton Co.): 305. Dixella vespertina. SOUTH DAKOTA Rapid Creek (Pennington Co.): 272. Dixa pullogruma. TEXAS Belfrage: 280. Dixa venosa. VIRGINIA Dead Run (Fairfax Co.): 265. Dixa inextricata. WASHINGTON Glacier (Whatcom Co.): 274. Dixa rhathyme. Mt. Rainier National Park, Longmire Springs: 256. Dixa arge; 285. Dix- ella thones. Puget: 268. Dixa melanderi. Peters: Topotypic North American Dixinae 7 REFERENCES CITED The citations of author, date and page in the List of Species refer to the bibliography in 'A Catalog of the Diptera of America North of Mexico" (U.S. D.A. Agricultural Handbook No. 276, 1965). Belkin, J.N., R.X. Schick and S.J. Heinemann 1966. Mosquito Studies (Diptera, Culicidae). VI. Mosquitoes originally de- scribed from North America. Amer. Entomol. Inst., Contrib. 1(6). oo Dp. Peters, T.M. 1966. [Descriptions of new species] In Peters, T.M. and E. F. Cook, 1966 (see below). Peters, T.M. and E. F. Cook 1966. The Nearctic Dixidae (Diptera). lications 5(5):231-278. 23 Dec. Entomol. Soc. Amer., Misc. Pub- INDEX TO SPECIFIC NAMES The references are to the numbers preceding the species names in the Rist of Species. aliciae, 282 arge, 256 aurora, 283 blax, 257 brevis, 258 californica, 284 centralis, 299 chalonensis, 306 clavata, 286 cornuta, 287 deltoura, 292 distincta, 259 dorsalis, 293 dyari, 294 fluvica, 260 fraterna, 261 fusca, 262 garretti, 275 hegemonica, 264 indiana, 295 inextricata, 265 johannseni, 266 lobata, 267 marginata, 296 melanderi, 268 modesta, 269 montana, 275 mystica, 263 neoaliciae, 297 neohegemonica, 270 nocheles, 300 notata, 271 nova, 298 occidentalis, 301 parva, 276 plexipus, 288 pullogruma, 272 repanda, 273 recens, 307 rhathyme, 274 rudis, 277 serrata, 302 similis, 278 simplex, 289 somnolenta, 290 spiralis, 291 techana, 303 terna, 279 thones, 285 universitatis, 304 venosa, 280 vespertina, 305 xavia, 281 £ eeatine — yet Pyar stti MOSQUITO STUDIES (Diptera, Culicidae) XI. MOSQUITOES ORIGINALLY DESCRIBED FROM ARGENTINA, BOLIVIA, CHILE, PARAGUAY, PERU, AND URUGUAY! By John N. Belkin, Robert X. Schick and Sandra J. Heinemann? This is the fourth in a series of papers providing information on the source of the original type material of mosquitoes described from the Americas (Bel- kin, Schick and Heinemann, 1965,1966; Peters, 1968). The concluding paper on species described from Brazil and unspecified localities in South America is in preparation. For explanation of the arrangement and method of presentation, the first paper of the series should be consulted. We are most grateful to Osvaldo H. Casal of the Instituto Nacional de Mi- crobiologfa, Buenos Aires, for his careful study of the type material in Argen- tina and for allowing us to publish his lectotype designations in this paper. We also thank Alan Stone for information on material contained in the U.S. Nation- al Museum. ARGENTINA List of Species 1. Anopheles (A.) annulipalpis Lynch Arribalzaga, 1878. TYPE: ¢, Bara- dero (Buenos Aires), Apr 1878 (NE, according to O.H. Casal). BIONOMICS: [Larvae in semipermanent turbid ground water with scanty vegetation. | 2. Anopheles (A.) holmbergi Del Ponte & Heredia, 1945 [=fluminensis]. TYPE: Holotype & (378) with genitalia slide (2293), Corpus, San Ignacio (Mis- iones), May 1944, R. L. Heredia (INM). BIONOMICS: [Larvae probably in per- manent or semipermanent ground waters. | *3. Anopheles (A.) argentinus (Bréthes, 1912) [=pseudopunctipennis]. TYPE: Lectotype 0, bearing label by Brethes, /Proterorhynchus argentinus Brethes, type, Tucumadn/ (BA; PRESENT DESIGNATION by O.H. Casal). BIONOMICS: [Larvae in small streams in mountains and foothills; water fresh, with mode- rate flow, and containing algae. | 1 Contribution from project ''Mosquitoes of Middle America’' supported by Public Health Service Research Grant AI-04379 and U.S. Army Medical Re- search and Development Command Research Contract DA-49-193-MD-2478. “Department of Zoology, University of California, Los Angeles, Califor- nia 90024. 10 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 4. Anopheles (A.) tucumanus Lahille, 1912 [=pseudopunctipennis|. TYPE: Holotype 2, Rio Sali (Tucuman), Dinelli (NE, according to O.H. Casal). BIO- NOMICS: [As for 3. argentinus. | 5. Anopheles (A.) patersoni Alvarado & Heredia, 1947 [=ssp. of pseudo- punctipennis|. TYPE: Eggs, Tucuman (probably NE, according to O. H. Casal). BIONOMICS: [As for 3. argentinus. | *6. Anopheles (Nyssorhynchus) albitarsis Lynch Arribalzaga, 1878. TYPE: ?, Baradero (Buenos Aires) [NE; original type material lost; redescription by Umana, Heredia and Siquot (1960:609-618) from type locality does not constitute a neotype designation, this material also lost]. BIONOMICS: [Larvae usually in mats of aquatic vegetation in large ponds, marshes and lagoons of overflow- ing rivers, and in large sunlit ground pools with vegetation. | 7. Anopheles (N.) rooti (Brethes, 1926) [=argyritarsis]. TYPE: o, Tu- cuman, N.C. Davis; only remaining material genitalia slide in poor condition, without sidepiece, with Bréthes label, ''Cellia Rooti Bréthes=argyritarsis auct. (nec R.D.), Tucuman, N.C. Davis dedit, III. 1956'"' (BA). BIONOMICS: [Larvae in all types of sunlit and partially shaded ground waters.| 8. Anopheles (N.) evansae (Bréthes, 1926). TYPE: Lectotype 0 genitalia slide found in INM (returned to BA), with following Brethes label, ''Cellia ev- ansi Brethes=tarsimaculata auct. (nec Goeld), Tucuman, N.C. Davis ded., III. 1926, Pr. micr. An.M.," in Del Ponte's hand '384" "DP14,'"" 'DP14= ? noroes- tensis G. y L., 1938, [signature in ink], Tipo de evansi [signature in black pen- cil] (BA; PRESENT DESIGNATION of O.H. Casal). BIONOMICS: [Larvae in marshes and marshy margins of clear freshwater pools, streams and lakes.| 9. Anopheles (N.) clarki Komp, 1943 [=noroestensis]|. TYPE: Holotype o& with genitalia slide, Monteros (Tucuman), June 1940, C.A. Alvarado (USNM, 56476; apparently lost, not mentioned in Stone and Knight, 1956b). BIONOM- ICS: [Larvae probably in shaded fresh water in wooded swamps, pools or stag- nant streams.| *10. Anopheles (N.) bachmanni Petrocchi, 1925 [=ssp. of triannulatus]. TYPE: o, 2, Corrientes, Entre Rios, Formosa (NE, according to O. H. Casal). BIONOMICS: [Larvae probably in freshwater pools and lakes, and along river margins, associated with vegetation. | 11. Anopheles (N.) perezi Shannon & Del Ponte, 1928 [=triannulatus bach- manni]. TYPE: 9, [Finca] Santa Barbara, Departamento de la Capital (Tucu-_ man) (NE, according to O.H. Casal). BIONOMICS: Larvae in a lake. 12. Anopheles (N.) davisi Paterson & Shannon, 1927 [=triannulatus bach- manni]. TYPE: o, @, larvae, Tres Pozos, [near] Embarcacion (Salta), 19-21 Apr 1927, Paterson, Shannon and Shannon (USNM, see Stone and Knight, 1956b: 277; INM, 1 2 with Shannon's labels, ''House, Embarcacion, Salta, 21.4. 27, R.C. Shannon/Anopheles (Nyssorhynchus) davisi Pat. & Snn (Cotype)'"'). BIO- NOMICS: Larvae in very large masses of green algae in a lake. 13. Toxorhynchites (Lynchiella) cavalierii Garcia & Casal. TYPE: Holo- type 2 with associated larval and pupal skins (c94), Puerto Iguazu about 50 m from the Hosteria Ruffino (Igu 15) (Misiones), 25 June 1965, Casal, Hepper and Garcia (INM). BIONOMICS: Larvae in a bromeliad 1 m above the ground. 14. Toxorhynchites (L.) tucumanus (Brethes, 1926) [=guadeloupensis]. TYPE: Lectotype & (1 of 6 with type labels) with following Brethes labels, ''Tu- cuman, 10-VII-926, E.G. Cabarrou/elevé chez moi/Megarhinus tucumanus Brethes" (BA; PRESENT DESIGNATION of O.H. Casal). BIONOMICS: [Lar- vae in leaf axils of bromeliads. | 15. Toxorhynchites (L.) arborealis (Shannon & Del Ponte, 1928) [=guade- Belkin et al: Topotypic South American Mosquitoes i loupensis]. TYPE: 9, Lules (Tucuman), Shannon and Del Ponte, 22 Mar 1927 [NE, according to O.H. Casal]. BIONOMICS: Larvae in a treehole. 16. Toxorhynchites (L.) separatus (Lynch Arribalzaga, 1891) [=ssp. of haemorrhoidalis}|. TYPE: Lectotype “ with Lynch label ''Megarhina separata n.sp.'’ and Del Ponte label ''Megarhinus haemorroidalis Fabr., D. P., X-49," according to original description collected in Formosa by E. L. Holmberg (BA; PRESENT DESIGNATION by O.H. Casal). BIONOMICS: [Larvae probably in leaf axils of bromeliads. | 17. Toxorhynchites (L.) lynchi (Dyar & Knab, 1906) [=haemorrhoidalis sep- aratus|. TYPE: Lectotype 0, only specimen in collection with Lynch's label, "Megarhyna haemorroidalis Fabr.,'' also with Del Ponte's label, 'Megarrhynus lynchi tipo, D. P. X-49,"' according to original description collected in Formo- sa by E.L. Holmberg (BA; PRESENT DESIGNATION by O.H. Casal). BIO- NOMICS: [As for 16. haemorrhoidalis separatus.| 18. Trichoprosopon (Runchomyia) paranensis (Bréthes, 1910) [Distinct spe- cies, according to O.H. Casal]. TYPE: Lectotype 9, the better of the 2 speci- mens bearing the following label, ''Tuyuparé, 25.1.1908, J.B. /Lynchiaria pa- ranensis" (BA; PRESENT DESIGNATION by O.H. Casal). BIONOMICS: [Lar- vae probably in leaf axils of cattails, Typha.| 19. Wyeomyia (Nunezia) lateralis Petrocchi, 1927. TYPE: 92, Zapla (Ju- juy), 16 Mar 1916, A. Neiva (NE, according to O.H. Casal). BIONOMICS: [Lar- vae probably in leaf axils of bromeliads.| *20. Wyeomyia (Davismyia) petrocchiae (Shannon & Del Ponte, 1928). TYPE: Holotype 2, Raco (Tucuman), 13 Feb 1927, Shannon and Del Ponte (USNM; see Stone and Knight, 1957b:126). BIONOMICS: [Larvae probably in treeholes.| 21. Wyeomyia (Davismyia) monoleua (Martini, 1931)[=petrocchiae]. TYPE: Holotype ¢, San Jose (Formosa), Oct 1925 (SMNS). BIONOMICS: [Larvae prob- ably in treeholes.| 22. Wyeomyia (Menolepis) leontiniae (Bréthes, 1910) [=leucostigma]. TYPE: Lectotype 2, the best of the 5 9, all without type label and all bearing Bréthes labels, ''Tuyupare, 25-I-1908, J.B" and ''Limatus Leontiniae" (BA; PRESENT DESIGNATION of O.H. Casal). BIONOMICS: [Larvae probably in leaf axils of cattails, Typha.| 23. Wyeomyia (Dendromyia) belkini Casal & Garcia, 1966. TYPE: Holo- type & with genitalia, larval and pupal skins (C57), intersection of the arroyo Ibicuy and route 101, about 20 km west of Cataratas (Igu 26) (Misiones), 26 June 1965, Hepper, Garcia and Casal (INM). BIONOMICS: Adults taken in the forest interior during the day, probably attracted to a human host. Larvae in bamboo internodes. 24. Wyeomyia (D.) typharum (Shannon & Del Ponte, 1928) [=melanocepha- la]. TYPE: Lectotype 9, bearing label probably in Del Ponte's hand 'Dendro- myia typharum" and also with following labels (present on other @), "Jujuy, Ledesma, 3.19.26/Davis and Shannon/49036" and ''Metatypus"” (BA; PRESENT DESIGNATION of O.H. Casal). BIONOMICS: Larvae in leaf axils of cattails, Typha. 25. Phoniomyia muehlensi (Petrocchi, 1927). TYPE: Holotype, sex un- known, Chaco, Oct-Nov 1924, J. Petrocchi (NE, according to O. H. Casal). BI- ONOMICS: [Larvae in leaf axils of bromeliads.]| 26. Limatus exhibitor Shannon & Del Ponte, 1928 [=durhamii]. TYPE: Lectotype & with following label, "Iguazu, Mis.[iones] 18.6.27, Shannon & Del Ponte" and in Shannon's hand, ''Limatus exhibitor Snn & D. P’’ (INM; PRESENT DESIGNATION of O. H. Casal). BIONOMICS: [Larvae in broken bamboo and in artificial containers at the type locality (O. H. Casal).| £2 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 . 27. Sabethes (S.) neivai Petrocchi, 1927 [=albiprivus]|. TYPE: Holotype 9, Santa Clara (Jujuy), May 1916, Arturo Neiva (NE, according to O.H. Casal). BIONOMICS: [Larvae probably in treeholes or bamboo internodes. | 28. Coquillettidia (Rhynchotaenia) fasciolata (Lynch Arribalzaga, 1891). TYPE: Originally described from 2 from Navarro (Buenos Aires), Mar 1886, F. Lynch (NE), neotype ?, Parque Lezama, Capital Federal (Buenos Aires), 13 Dec 1928, Carillo (INM; designation of Castro and Bressanello, 1952:232). BI- ONOMICS: [Larvae probably on rootlets of grasses and herbaceous vegetation in mud and fine sediment in very shallow water on the margins of swamps, ponds or streams. | 29. Coquillettidia (Rhynchotaenia) araozi (Shannon & Del Ponte, 1928) [= venezuelensis]|. TYPE: ?, Ledesma (Jujuy), 20 Mar 1926, Davis and Shannon; no specimens of type series in BA, according to O. H. Casal (USNM; lectotype to be designated by Alan Stone). BIONOMICS: [Larvae probably as for 28. fas- ciolata.| ¢ taken on animal bait. 30. Uranotaenia lanei Martinez & Prosen, 1953. TYPE: Holotype ¢, For- mosa (BA, according to O.H. Casal). BIONOMICS: [Larvae probably in per- manent or semipermanent ground waters. | 31. Uranotaenia monilis Shannon & Del Ponte, 1928. TYPE: Lectotype ? bearing the labels "Resistencia, Chaco, 20.2.27, R.C. Shannon/Uranotaenia monilis Snn. & D.P.' in Shannon's hand (INM; PRESENT DESIGNATION by O.H. Casal). BIONOMICS: [Larvae probably in large lowland swamps or in other sunlit ground waters with some aquatic vegetation. | *32. Uranotaenia nataliae Lynch Arribalzaga, 1891. TYPE: Lectotype adult, only the thorax and first abdominal segment remaining, with a label by Lynch "Uranotaenia nataliae n.sp., Tigre" [at the confluence of the Rio Las Conchas and Rio Parana (Buenos Aires), Enrique Lynch, brother of Félix] (BA; PRES- ENT DESIGNATION by O.H. Casal). BIONOMICS: [Larvae probably in large open swamps with abundant aquatic vegetation such as Pistia and sedges. | 33. Uranotaenia capitis Shannon & Del Ponte, 1928. TYPE: Holotype 2, Ledesma (Jujuy), 20 Mar 1926, Davis (USNM). BIONOMICS: [As for 32. natal- iae.| *34, Uranotaenia pulcherrima Lynch Arribalzaga, 1891. TYPE: , [prob- ably] bank of Rio Lujan (Buenos Aires), E. L. Holmberg (NE, according to O. H. Casal). BIONOMICS: [Larvae probably in swamps, stream margins or other permanent ground waters. | : | 35. Uranotaenia urania Shannon & Del Ponte, 1928 [=pulcherrima]. TYPE: Lectotype o (2355) with genitalia slide, Resistencia (Chaco), 20 Feb 1927, Shan- non and Del Ponte (USNM, designation of Stone and Knight, 1957c:200). BIONOM- ICS: [As for 34. pulcherrima.| 36. Uranotaenia elnora Paterson & Shannon, 1927 [ssp. of pulcherrima]. TYPE: Holotype 2, Tres Pozos near Embarcacion (Salta), 19 Apr 1927, Elnora S. Shannon (USNM). BIONOMICS: [Probably as for 34. pulcherrima. | *37. Aedeomyia squamipennis (Lynch Arribalzaga, 1878). TYPE: o, ¢, Ba- radero (Buenos Aires), Apr 1878, F. Lynch (NE, according to O.H. Casal). BIONOMICS: [Larvae in lakes, ponds and permanent ground pools with abundant vegetation and algae. | 38. Psorophora (P.) lynchi Brethes, 1916 [=ciliata]. TYPE: Lectotype 9, the best of the 4 female syntypes in the BA, and in good condition, bearing the labels ''B.[uenos] Aires, 20.11.1916, J.B. /Psorophora Lynchi Br." (the other 3 females are similarly labeled) (BA; PRESENT DESIGNATION by O.H. Casal). BIONOMICS: [Larvae probably in unshaded temporary rain pools. | Belkin et al: Topotypic South American Mosquitoes 13 ‘39. Psorophora (P.) holmbergii Lynch Arribalzaga, 1891. TYPE: Lecto- type 2, the one specimen in the collection of the BA bearing the following label by F. Lynch, 'Psorophora Holmbergi n.sp., Chaco'' [Formosa], E.L. Holm- berg (BA; PRESENT DESIGNATION by O.H. Casal). BIONOMICS: [Larvae in temporary rain pools.| 40. Psorophora (P.) agoggylia Dyar, 1922 [=holmbergii]. TYPE: Lecto- type 2, Rio Tapenaga, Colonia Florencia, Gran Chaco (Formosa [Santa Fe)]), 1903, E.R. Wagner (USNM, 25756; designation of Stone and Knight, 1955:286). BIONOMICS: [Larvae probably in temporary rain pools.| 41. Psorophora (P.) stigmatephora Dyar, 1922 [=pallescens]. TYPE: Lec- totype & (1660) with genitalia slide, Rio Tapenga, Colonia Florencia, Gran Chaco (Formosa [Santa Fe]), 1903, E.R. Wagner (USNM, 25756; designation by Stone and Knight, 1955:286). BIONOMICS: [Larvae probably in temporary rain pools. |] 42. Psorophora (Janthinosoma) centrale Brethes, 1910 [=ferox]. TYPE: Lectotype 2 with the following labels, "Buenos Aires, 4. XII. 1907, B./Janthino- soma centralis [sic] Bréthes'' in Brethes' hand, type locality cited as "Islas del Parana, Buenos Aires" in the original description (BA; PRESENT DESIGNA- TION by O. H. Casal). BIONOMICS: [Larvae probably in temporary rain and floodwater pools in wooded areas.| 43. Psorophora (J.) chaquensis Paterson & Shannon, 1927 [=lutzii]. TYPE: Holotype °, Tres Pozos, near Embarcacion (Salta), 20 Apr 1927, probably Paterson and Shannon (NE, according to Casal). BIONOMICS: [Larvae prob- ably in temporary rain and floodwater pools in wooded areas.| 44. Psorophora (J.) bruchi Petrocchi, 1927 [=varipes|]. TYPE: 0, 2, Pro- vincia de Buenos Aires, 1 Apr 1925, C. Bruch (NE, according to O. H. Casal). BIONOMICS: [Larvae probably in temporary rain and floodwater pools in wooded areas. | *45. Psorophora (Grabhamia) confinnis Lynch Arribalzaga, 1891. TYPE: Lectotype 2, the only female of the original 5 specimens remaining in the BA, in fairly good condition, with the following label by F. Lynch, ''Taeniorhynchus confinnis F. Lch., Chaco [Formosa],'' E.L. Holmberg (BA; PRESENT DESIG- NATION by O.H. Casal). BIONOMICS: [Larvae probably in open temporary ground pools of all types, especially hoofprints and road ruts.]| 46. Psorophora (G.) paulli (Paterson & Shannon, 1927). TYPE: Lectotype 2, proboscis broken, 2 legs missing and abdomen glued on a card, with the fol- lowing labels in Shannon's handwriting, ''Tres Pozos, Salta, 20.4.27, Shannon & Snn/Types Psorophora Paulli Paterson & Shannon" (INM; PRESENT DESIG- NATION by O.H. Casal). BIONOMICS: [Larvae probably in temporary ground pools or overflows of ponds or streams. | 47. Aedes (O.) meprai Martinez & Prosen, 1953 [=angustivittatus]|. TYPE: Holotype 2, Reserva Nacional ''Finca del Rey,’’ Departamento de Anta (Salta), Nov 1952, Martinez and Prosen (BA). BIONOMICS: [Larvae in temporary rain pools.] Adults captured on human bait in daytime in forest. 48. Aedes (Ochlerotatus) lynchii (Bréthes, 1910) [=crinifer]. TYPE: Lec- totype 2, the best of 9 females, in fairly good condition, all from Buenos Aires with the following labels, "6139/Buenos Aires, J. Brethes 19.1. 1903/Culex Lynchii Br.,'' the latter two by Brethes (BA; PRESENT DESIGNATION by O. H. Casal). BIONOMICS: [Larvae probably in temporary rain pools. | 49. Aedes (O.) tapinops (Brethes, 1917) [=crinifer]. TYPE: Lectotype c, in fairly good condition, with the following labels by Brethes, ''S. Isidro [Buenos Aires], II.2.1917. J. B./Culex tapinops Bret.'' (BA; PRESENT DESIGNATION 14 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 by O.H. Casal). BIONOMICS: [Larvae probably in temporary rain pools. | 50. Aedes (O.) iguazu Shannon & Del Ponte, 1928 [=crinifer]|. TYPE: o, 9 [Cataratas del Iguazu], Misiones (3 2, without type labels but collected at Igua- zu Falls by Shannon and Shannon in Oct 1927, are present in the USNM; no specimens of the type series are in the INM). BIONOMICS: [Larvae probably in temporary rain pools.| 51. Aedes (O.) araozi Shannon & Del Ponte, 1928 [=milleri]. TYPE: Lec- totype co’, in fairly good condition, with the following labels in Shannon's hand, "Quebrada San Lorenzo, Salta, 4. VIII.27, R.C. Shannon/A. 154/Aedes araozi Shannon & Del Ponte," with genitalia slide (655) (INM; PRESENT DESIGNATION by O. H. Casal). BIONOMICS: Larvae in a small rockhole covered with dead leaves very near a small torrential stream. 52. Aedes (O.) patersoni Shannon & Del Ponte, 1928. TYPE: Lectotype §, in very bad condition, a mesothoracic leg and an antenna are missing and the mesonotum is largely rubbed, with the following labels, ''San Pedro de Jujuy, 27.4.26/Inst. Bac. Ent. nota 44/Shannon and Shannon" (INM; PRESENT DES- IGNATION by O.H. Casal; the same specimen improperly designated as Holo- type by Garcia & Ronderos, 1963:32). BIONOMICS: Larvae in a temporary swampy area formed by rains. 53. Aedes (O.) raymondi Del Ponte, Castro & Garcia, 1951. TYPE: Holo- type @ (587), San Pedro (Jujuy), 27 Apr 1926, Shannon and Shannon (INM). BI- ONOMICS: [Larvae probably in temporary rain pools. | *54. Aedes (O.) confirmatus (Lynch Arribalzaga, 1891) [=scapularis|. TYPE: Lectotype @, in good condition, with the following labels by F. Lynch, "Ochle- rotatus cofirmatus [sic], F Lynch, Navarro [Buenos Aires],' May 1887 (BA; PRESENT DESIGNATION by O. H. Casal). BIONOMICS: [Larvae in temporary rain pools.| 55. Haemagogus (Stegoconops) petrocchiae Martinez, Carcavallo & Prosen, 1961 (=ssp. of capricornii]. TYPE: Holotype o, Salvador Mazza (Pocitos), Departamento General San Martin (Salta), Jan 1960, Martinez and Carcavallo (BA). BIONOMICS: Females taken biting on human bait at dusk and at night in woods without artificial lights. [Larvae-probably in treeholes. | 56. Haemagogus (Stegoconops) spegazzinii Brethes, 1912. TYPE: Holotype ? [with the following labels, ''No 727 3/Jujuy/ Haemagogus spegazzinii Brethes" and 2 slides (3028), one with a wing and the other with genitalia, an antenna and a palpus], Jujuy, 1907, D. Carlos Spegazzini(BA). BIONOMICS: [Larvae prob- ably in treeholes.] Adult(s) taken in deep woods. 57. Haemagogus (Stegoconops) uriartei Shannon & Del Ponte [=spegazzinii]. TYPE: Holotype % (128-3) with genitalia slide (2353) and pupal skin slide (V 3), Vipos (Tucuman), 22 Mar 1927, Shannon and Del Ponte (USNM; see Stone and Knight, 1955:289). BIONOMICS: Holotype bred from larva in a treehole. 58. Culex (Lutzia) patersoni Shannon & Del Ponte, 1928 [=bigoti]. TYPE: Lectotype & (2356) with genitalia slide, San Pedro (Jujuy), 27 Apr 1926, Shan- non and Shannon (USNM; designation of Stone and Knight, 1957a:54). BIONOMICS: [Larvae in ground pools, rockholes and artificial containers. | 59. Culex (C.) ameliae Casal, 1967. TYPE: Holotype o, with associated larval and pupal skins (Ga 137-31), San Pedro-El Dorado road, about 10 km from San Pedro (Misiones), 11 Mar 1966, Garcia and Casal (INM). BIONOM- ICS: Larvae in an extensive flooded depression in the middle of the forest crossed by the road. 60. Culex (C.) interfor Dyar, 1928 [=bidens]. TYPE: Lectotype & (532) with genitalia slide (2364), in train between Tucuman and Jujuy (Jujuy, Salta or Belkin et al: Topotypic South American Mosquitoes 15 Tucuman), 4 May 1927, M. Kisliuk Jr. (USNM; designation by Stone and Knight 1957a:51). BIONOMICS: [Larvae probably in permanent and semipermanent ground waters.| Adults taken at light on train. 61. Culex (C.) brethesi Dyar, 1919. TYPE: Holotype “ (on slide with the genitalia and a female) [San Isidro (Buenos Aires), Apr or May 1916] (BA). BI- ONOMICS: [Larvae probably in permanent or semipermanent ground waters. | 62. Culex (C.) castroi Casal & Garcia, 1967. TYPE: Holotype & with as- sociated larval and pupal skins (Ga 289-11), Canal 6 and Parana de las Palmas, Estacion Agropecuaria Delta del I.N. T. A. (near Otamendi) (Buenos Aires), 27 July 1966, Garcia and Casal (INM). BIONOMICS: Larvae in leaf axils of Eryn- sium sp. *63. Culex (C.) dolosus (Lynch Arribalzaga, 1891). TYPE: Lectotype o, in very bad condition, partially covered by fungi, the head and the posterior ab- dominal segments missing, with the following label by F. Lynch, ''Heterony- cha dolosa n.sp.,'' no locality label, the following 5 localities given in the orig- inal description: Las Conchas, Zarate, Baradero, Navarro and Chacabuco (all Buenos Aires) (BA; PRESENT DESIGNATION by O.H. Casal). BIONOMICS: [Larvae in permanent and semipermanent ground waters. | 64. Culex (C.) bonariensis Bréthes, 1916 [=dolosus]. TYPE: Lectotype, with type label but without locality label, and genitalia slide (V) [San Isidro (Buenos Aires)|, May (BA; PRESENT DESIGNATION by O.H. Casal). BIO- NOMICS: [Probably as for 63. dolosus. | 65. Culex (C.) eduardoi Casal & Garcia, 1968. TYPE: Holotype with as- sociated larval and pupal skins (Ga 314-17), Canal 6 and Parana de las Palmas, Estacion Agropecuaria Delta de I.N. T.A. (near Otamendi) (Buenos Aires), 10 Aug 1966, Garcia and Casal (INM). BIONOMICS: Larvae in a marshy depres- sion. 66. Culex (C.) fernandezi Casal, Garcia and Cavalieri, 1966. TYPE: Holo- type o with associated larval and pupal skins (Ga 85-15), stream in Piquirenda (Salta), 26 Nov 1965, Fernandez, Hepper, Garcia and Casal (INM). BIONOM- ICS: Larvae in axils of epiphytic or terrestrial bromeliads. 67. Culex (C.) hepperi Casal & Garcia, 1967. TYPE: Holotype & with as- sociated larval and pupal skins (Ga 268-10), Canal 6 and Parana de las Palmas, Estacion Agropecuaria Delta de I.N. T. A. (near Otamendi) (Buenos Aires), 30 Apr 1966, Garcia and Casal (INM). BIONOMICS: Larvae in leaf axils of Eryn- gium. 68. Culex (C.) lahillei Bachmann & Casal, 1962. TYPE: Holotype & (1124) with genitalia slide (5219), Achiras (Cordoba), Mar 1938, Del Ponte (INM). BI- ONOMICS: [Larvae probably in permanent and semipermanent ground waters. | 69. Culex (C.) maxi Dyar, 1928. TYPE: Lectotype o on slide 2360 and some legs on pin mount, San Pedro (Jujuy), 11 June 1927, M. Kisliuk Jr. (USNM; designation by Stone and Knight, 1957a:53). BIONOMICS: [Larvae in permanent and semipermanent ground waters. | 70. Culex (C.) autumnalis Weyenbergh, 1882 [=pipiens quinquefasciatus]. TYPE: o, °, larva, pupa, Rio Primero (Cérdoba) (? Universidad de Cordoba). BIONOMICS: [Larvae in large artificial containers and contaminated ground water. | 71. Culex (C.) saltanensis Dyar, 1928. TYPE: Holotype o’, Campo Santo de Salta (Salta), 12 May 1927, M. Kisliuk Jr. (USNM). BIONOMICS: [Larvae probably in permanent or semipermanent ground waters. | *72. Culex (Allimanta) tramazayguesi Duret, 1954. TYPE: Holotypeo" (M.R. 110, E.1), Monte Coman (Mendoza), 16 Dec 1953, Bejarano and Duret (CMPHM). 16 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 BIONOMICS: [Larvae found in hypersaline ground water near a thermal spring (Casal).]| 73. Culex (Melanoconion) aliciae Duret, 1953. TYPE: Holotype o (Mis. 13, E 8) with genitalia slide, Cerro Azul, Arroyo San Juan (Misiones), 2 May 1949, Bejarano and Duret (CMPHM). BIONOMICS: [Larvae probably in stream pools or other permanent or semipermanent ground waters. | 74. Culex (Mel.) bejaranoi Duret, 1953. TYPE: Holotype o (A. U. 93, E. 20) with genitalia slide, Bernardo de Irigoyen (Misiones), 18 Mar 1951, Duret (CMPHM). BIONOMICS: [Larvae found in large ground pool in a probably flooded forest (Casal). | 75. Culex (Mel.) intrincatus Brethes, 1916. TYPE: Holotype <, lowlands of San Isidro (Buenos Aires), 15 May 1916, Brethes (BA). BIONOMICS: [Lar- vae probably in permanent or semipermanent ground waters.| 76. Culex (Mel.) martinezi Casal & Garcia, 1968. TYPE: Holotype with associated larval and pupal skins (Ga 418-17), km 34 of the Vespucio-San Pedri- to road, near Vespucio (Salta), 6 Oct 1966, Garcia, Hepper and Fernandez (INM). BIONOMICS: Larvae taken at the margins of an arroyo. 77. Culex (Mel.) misionensis Duret, 1953. TYPE: Holotype & (A.U. 23, E. 14) with genitalia slide, Aristobulo del Valle (Misiones), 28 Feb 1951, Duret (CMPHM). BIONOMICS: [Larvae probably in permanent or semipermanent ground waters. | 78. Culex (Mel.) orfilai Duret, 1953. TYPE: Holotype & (D.O 69, E. 20) with genitalia slide, Iguazu, Villa Tacuara (Misiones), 21 Nov 1951, Duret (CMPHM). BIONOMICS: [Larvae probably in permanent or semipermanent ground waters. | 79. Culex (Mel.) pavlovskyi Casal & Garcia, 1967. TYPE: Holotype & with genitalia slide (660331), Aeropuerto de Camba Punta, near Corrientes (Corrien- tes), 4 Mar 1966, O.H. Casal (INM). BIONOMICS: Adults taken at a lighted window. [Larvae probably in permanent or semipermanent ground waters.| 80. Culex oblita (Lynch Arribalzaga, 1891) [NEW COMBINATION; NOMEN DUBIUM]. TYPE: Holotype “ [only a leg and a wing remain], Navarro, near Arroyo de las Saladas (Buenos Aires), Mar 1884[F. Lynch] (BA). 81. Sayomyia australis (Shannon & Del Ponte, 1928). TYPE: Lectotype §, in good condition, with the following labels, ''Ins. Bac. Ent. nota 195/Finca San- ta Barbara, 6.IV.27 Tuc.[uman, Departamento de la Capital]/R.C. Shannon coll & Del Ponte/Chaoborus australis Shannon & Del Ponte" (INM; PRESENT DESIGNATION by O. H. Casal). BIONOMICS: Adults collected at lights. [Lar- vae probably in permanent or semipermanent still ground waters. | | 82. Sayomyia elnorae (Shannon & Del Ponte, 1928). TYPE: 2, Tres Pozos, near Embarcacion (Salta), E.S. Shannon (LU). BIONOMICS: Adults collected at light near a lake. [Larvae probably in lakes and other permanent and semi- permanent still ground waters. | 83. Corethrella arborealis Shannon & Del Ponte, 1928 [=appendiculata]. TYPE: Holotype o [(174) and wing slide (389)], Vipos [Tucuman], 4 Mar 1927, E. Del Ponte (INM). BIONOMICS: Holotype bred from pupa from treehole. 84. Corethrella puella Shannon & Del Ponte, 1928. TYPE: Lectotype @, in good condition, with the following labels, ''Ledesma, Jujuy, 30.3.27/R.C. Shan- non coll./Corethrella puella S & DP," the last label by Del Ponte (INM; PRES- ENT DESIGNATION by O. H. Casal). BIONOMICS: Adults collected in bathroom. [Larvae probably in permanent and semipermanent ground water with dense vegetation. | 85. Corethrella quadrivittata Shannon & Del Ponte, 1928. TYPE: Lectotype Belkin et al: Topotypic South American Mosquitoes 17 o", in very good condition, with the following labels, ''Tres Pozos, Salta, 20.4. 27/Shannon & Shannon. /Corethrella quadrivittata Shannon y Del Ponte," the last label by Shannon (INM; PRESENT DESIGNATION by O. H. Casal). BIO- NOMICS: Females taken at light near a lake. Larvae in aquatic vegetation (Pis- tia and others) in lakes. 86. Lutzomiops davisi (Shannon & Del Ponte, 1928). TYPE: Lectotype 9, in fairly good condition, with the following labels, "Ins. Bac. Ent. nota 32-4/ Concepcion, Tuc.[umdan] 4.7.26/R.C. Shannon/390/Corethrella davisi Shannon & Del Ponte," the last label by Shannon (INM; PRESENT DESIGNATION by O. H. Casal). BIONOMICS: Larvae in a ground pool with semistagnant water and with vegetation at the margins. 87. Nothodixa atrovittata (Edwards, 1930). TYPE: Holotype “, Bariloche [San Carlos de Bariloche] (Rio Negro), elev. 2450 ft, 25 Oct-1 Dec 1926, F. W. Edwards (BM). BIONOMICS: [Larvae probably in marginal vegetation or flo- tage in small creeks. | 88. Nothodixa nitida (Edwards, 1930). TYPE: Holotype 0, Bariloche [San Carlos de Bariloche] (Rio Negro), elev. 2450 ft, 25 Oct-1 Dec 1926, F.W. Ed- wards (BM). BIONOMICS: [Larvae probably in marginal vegetation or flotage in small creeks.| 89. Dixella argentina (Alexander, 1920). TYPE: Holotype 0, La Granja, Alta Gracia (Cordoba), 1-8 Apr 1920, Charles Bruch(A). BIONOMICS: [Larvae probably in various types of permanent and semipermanent ground waters, es- pecially at margins of small streams. | List of Localities BUENOS AIRES [Locality not specified]: 38. Psorophora (P.) lynchi; 44. Psorophora (J.) bruchi; 48. Aedes (O.) lynchii. Baradero: 1. Anopheles (A.) annulipalpis; 6. Anopheles (N.) albitarsis; 37. Aedeomyia squamipennis; 63. Culex (C.) dolosus. Chacabuco: 63. Culex (C.) dolosus. Las Conchas: 63. Culex (C.) dolosus. Lujan (Rio): 34. Uranotaenia pulcherrima. Navarro: 28. Coquillettidia (R.) fasciolata; 54. Aedes (O.) confirmatus (banks of Rio Salado); 63. Culex (C.) dolosus. Otamendi (Ingeniero Romulo) (Canal 6 and Parana de las Palmas, Estacion Agropecuaria Delta de I. N. T. A.): 62. Culex (C.) castroi; 65. Culex (C.) eduar- doi; Culex (C.) hepperi. Parana, Islas del (Tuyuparé, a brook in the delta of the Parana): 18. Trich- oprosopon (R.) paranensis; 22. Wyeomyia (M.) leontiniae; 42. Psorophora (J.) centrale. San Isidro: 49. Aedes (O.) tapinops; 61. Culex (C.) brethesi; 64. Culex (C.) bonariensis; 75. Culex (Mel.) intrincatus (lowlands). Tigre: 32. Uranotaenia nataliae (Rio Las Conchas). Zarate: 63. Culex (C.) dolosus. CHACO | [Locality not specified]: 25. Phoniomyia mueblensi. Resistencia: 31. Uranotaenia monilis; 35. Uranotaenia urania. 18 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 CORDOBA Achiras: 68. Culex (C.) lahillei. La Granja, Alta Gracia: 89. Dixella wiventina. Primero (Rio): 70. Culex (C.) autumnalis. CORRIENTES [Locality not specified]: 10. Anopheles (N.) bachmanni. Corrientes: 79. Culex (Mel.) pavlovskyi (Aeropuerto de Camba Punta). ENTRE RIOS [Locality not specified]: 10. Anopheles (N.) bachmanni. FORMOSA [Locality not specified]: 10. Anopheles (N.) bachmanni; 16. Toxorhynchites (L.) separatus; 17. Toxorhynchites (L.) lynchi; 30. Uranotaenia lanei; 39. Psor- ophera (P.) holmbergii; 45. Psorophora (G.) confinnis. San Jose: 21. Wyeomyia (Dav.) monoleua. J UJUY [Locality not specified]: 56. Haemagogus (S.) spegazzinii; 60. Culex (C.) in- terfor (on train between Tucuman and Jujuy). Ledesma: 24. Wyeomyia (D.) typharum; 29. Coquillettidia (R.) araozi; 33. Uranotaenia capitis; 84. Corethrella puella (Cafiitas Viejo). San Pedro: 52. Aedes (O.) patersoni; 53. Aedes (O.) raymondi; 58. Culex (Lutzia) patersoni; 69. Culex (C.) maxi. — Santa Clara: 27. Sabethes (S.) neivai. Zapla: 19. Wyeomyia (N.) lateralis. MENDOZA Monte Coman: 72. Culex (All.) tramazayguesi. MISIONES [Locality not specified, probably Cataratas del Iguazu]: 50. Aedes (O.) iguazu. Aristobulo del Valle: 77. Culex (Mel.) misionensis. Bernardo de Irigoyen: 74. Culex (Mel.) bejaranoi. Cataratas: 23. Wyeomyia (D.) belkini (20 km west of). Cerro Azul, Arroyo San Juan: 73. Culex (Mel.) aliciae. Corpus, San Ignacio: 2. Anopheles (A.) holmbergi. Iguazu (Cataratas del): 26. Limatus exhibitor. Iguazu, Villa Tacuara: 78. Culex (Mel.) orfilai. Puerto Iguazu: 13. Toxorhynchites (L.) cavalierii (50m from Hosteria Ruf- fino). San Pedro: 59. Culex (C.) ameliae (10 km west of, on San Pedro-El Dorado road). RIO NEGRO San Carlos de Bariloche, 2450 ft: 87. Nothodixa atrovittata; 88. Nothodixa nitida. SALTA [Locality not specified]: 60. Culex (C.) interfor (on train between Tucuman and Jujuy). Belkin et al: Topotypic South American Mosquitoes 1? Campo Santo de Salta: 71. Culex (C.) saltanensis. Piquirenda: 66. Culex (C.) fernandezi. Reserva Nacional "Finca del Rey," Departamento de Anta: 47. Aedes (O.) meprai. Salvador Mazza (Pocitos), Departamento General San Martin: 55. Haema- ogus (S.) petrocchiae. San Lorenzo (Quebrada de), near Salta: 51. Aedes (O.) araozi. Tres Pozos, near Embarcacidn: 12. Anopheles (N.) davisi; 36. Uranotaenia ere elnora; 43. Psorophora (J.) chaquensis; 46. Psorophora (G.) paulli; 82. Sayomy- ia elnorae; 85. Corethrella quadrivittata. Vespucio: 76. Culex (Mel.) martinezi (km 34 of Vespucio-San Pedrito road). SANTA FE Tapenaga (Rio), Colonia Florencia, Gran Chaco: 40. Psorophora (P.) agog- gylia; 41. Psorophora (P.) stigmatephora. TUCUMAN [Locality not specified]: 3. Anopheles (A.) argentinus; 5. Anopheles (A.) patersoni; 7. Anopheles (N.) rooti; 8. Anopheles (N.) evansae; 14. Toxorhynchi- tes (L.) tucumanus; 60. Culex (C.) interfor (on train between Tucuman and Ju- juy). Concepcion: 86. Lutzomiops davisi (on road 5 km W). Lules: 15. Toxorhynchites (L.) arborealis. Monteros: 9. Anopheles (N.) clarki. Raco: 20. Wyeomyia (Dav.) petrocchiae. Salf (Rio): 4. Anopheles (A.) tucumanus. Santa Barbara (Finca), Departamento dela Capital: 11. Anopheles (N.) per- ezi; 81. Sayomyia australis. Vipos: 57. Haemagogus (S.) uriartei; 83. Corethrella arborealis. BOLIVIA List of Species 1. Anopheles (Kerteszia) boliviensis Theobald, 1905. TYPE: Holotype e, Songo [Zongo] (La Paz), date not specified, M. Bird (BM). BIONOMICS: |Lar- vae probably in leaf axils of bromeliads. | 2. Toxorhynchites (Ankylorhynchus) hexacis (Martini, 1931). TYPE: Holo- type 9, Yungas de Coroico, Nor Yungas (La Paz), elev. 1000 m (NE). BIONOM- ICS: [Larvae probably in treeholes or leaf axils.| 3. Trichoprosopon (Ctenogoeldia) magnum (Theobald, 1905). TYPE: Holo- type ?, San Antonio [de Mapiri] (La Paz), M. Biro (HNM). BIONOMICS: [Lar- vae probably in leaf axils and flower bracts of Calathea or other Marantaceae. | 4. Mansonia (M.) fonsecai (Pinto, 1932) [=indubitans]|. TYPE: Holotype ? (160), Los Naranjos [Naranjo] (Santa Cruz), 23 Feb 1925, O. da Fonseca (IOC). BIONOMICS: [Larvae probably attached to floating vegetation (Pistia) in ponds or lakes.| 5. Coquillettidia (Rhynchotaenia) nitens (Cerqueira, 1943). TYPE: Holo- type 2, Puerto Suarez, Chiquitos (Santa Cruz), July 1939 (IOC). BIONOMICS: [Larvae probably on rootlets of grasses and other herbaceous vegetation in mud or fine sediment in very shallow water on margins of swamps, ponds or lakes.| 20 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 6. Orthopodomyia bacigalupoi Martinez & Prosen, 1958 [=fascipes]. TYPE: Holotype o, Tact, Buenavista, Ichilo (Santa Cruz), Apr 1955, A. Martinez and A. F. Prosen (BA). BIONOMICS: Larvae in treeholes. 7. Psorophora (Janthinosoma) circumflava Cerqueira, 1943. TYPE: Holo- type °, Axiamas, Caupolican (La Paz), Dec 1941 (IOC). BIONOMICS: | Larvae probably in temporary ground pools or overflows of ponds or streams. | 8. Psorophora (J.) melanota Cerqueira, 1943. TYPE: Holotype <, Riber- alta, Terr. de Colonias (El Beni), Dec 1939 (IOC). BIONOMICS: [Larvae prob- ably in temporary ground pools or overflows of ponds or streams.| 9. Psorophora (Grabhamia) chiquitana Pinto, 1932 [=confinnis]. TYPE: Holotype 2 (163), Los Naranjos [Naranjo] (Santa Cruz), 1925, O. da Fonseca (IOC). BIONOMICS: [Larvae probably in temporary ground pools or overflows of ponds or streams.| 10. Psorophora (G.) dimidiata Cerqueira, 1943. TYPE: Holotype“, La- eunillas, Valle Grande (Santa Cruz), Feb 1940 (IOC). BIONOMICS: [Larvae probably in temporary ground pools or overflows of ponds or streams. | 11. Aedes (Ochlerotatus) bejaranoi Martinez, Carcavallo & Prosen, 1960. TYPE: Holotype o°, Yungas del Palmar, between km 90 and 110 on road from Cochabamba to Villa Tunari [San Antonio), Chapare (Cochabamba), elev. 3000- 3900 m, Feb 1959, A. Martinez (BA). BIONOMICS: Larvae in pools of very cold, slowly flowing water and with dense vegetation and abundant filamentous algae. Adults of both sexes resting in vegetation on margins of pools. 12. Aedes (O.) oroecetor Martini, 1931 miter TYPE: Lectotype with associated genitalia slide, Sorata (La Paz), elev. 2300 m (BM; designation of Mattingly, 1955:29). BIONOMICS: [Larvae probably in shaded stream bed pools with organic debris and surrounded by vegetation. | 13. Aedes (Howardina) aurivittatus Cerqueira, 1943. TYPE: Holotype 9, Pampa Grande, Florida (Santa Cruz), Feb 1940 (IOC). BIONOMICS: [Larvae probably in leaf axils of bromeliads, possibly in treeholes.| 14. Aedes (H.) delpontei Martinez & Prosen, 1955. TYPE: Holotype , be- tween km 125 (Limbo, elev. 2000 m) and km 145 (Locotal, elev. 1200 m) on road from Cochabamba to Villa Tunari [San Antonio], Chapare (Cochabamba), Jan 1949, Feb 1952 or Nov 1953, A. Martinez and A. F. Prosen (BA). BIONOMICS: [Larvae probably in leaf axils of bromeliads, possibly in treeholes. | 15. Aedes (H.) vanemdeni Martini, 1931. TYPE: Lectotype o, Yungas de Coroico, Nor Yungas (La Paz), elev. 1000 m (BM; designation of Mattingly, 1955:31). BIONOMICS: [Larvae probably in leaf axils of bromeliads, possibly in treeholes.| 16. Haemagogus (Stegoconops) lindneri Martini, 1931 [=speggazzinii or uri- artei|. TYPE: Holotype ?, [?San José de] Chiquitos (Santa Cruz), Oct 1926 1926 (SMNS). BIONOMICS: [Larvae probably in treeholes. | 17. Culex (C.) alticola Martini, 1931. TYPE: Lectotype 0, Serata [Sorata] (La Paz), elev. 2300 m, 19 Dec 1902 (BM; designation of Mattingly, 1955:31). BIONOMICS: [Larvae probably in permanent or semipermanent ground waters. | 18. Culex (C.) bidens Dyar, 1922. TYPE: Lectotype & with genitalia slide (1687), Rosario, near Lago Rogagua (El Beni), Nov 1921, W.M. Mann (USNM, 25760; designation of Stone and Knight, 1957a:47). BIONOMICS: [Larvae prob- ably in permanent and semipermanent ground waters.| Adults probably collec- ted in forest islands in generally open country. 19. Edwardsops boliviensis Lane & Heredia, 1956. TYPE: Holotype ¢, El Paylon [E1 Pailon| (Santa Cruz), 5-6 Feb 1955, Travassos, Barros and Albu- querque (FH). BIONOMICS: [Larvae probably in permanent or semipermanent ground pools, possibly in treeholes. | Belkin et al: Topotypic South American Mosquitoes 21 List of Localities BENI (EL) Riberalta: 8. Psorophora (J.) melanota. Rosario, near Lago Rogaguaz 18. Culex (C.) bidens. San Antonio: (See under LA PAZ). COCHABAMBA | Chapare Prov.; road from Cochabamba to Villa Tunari (San Antonio): 11. Aedes (O.) bejaranoi (Yungas del Palmar, between km 90 and 110; elev. 3000- 3900 m); 14. Aedes (H.) delpontei (between km 125, elev. 2000 m, and km 145, elev. 1200 m). : LA PAZ Axiamas, Caupolican Prov.: 7. Psorophora (J.) circumflava. Coroico (Yungas de) elev. 1000 m, Nor Yungas Prov.: 2. Toxorhynchites (A.) hexacis; 15. Aedes (H.) vanemdeni. San Antonio de Mapiri: 3. Trichoprosopon (Ctenogoeldia) magnum. Sorata: 12. Aedes (O.) oroecetor; 17. Culex (C.) alticola. Zongo (Songo): 1. Anopheles (K.) boliviensis. SANTA CRUZ Buenavista (Tact), Ichilo Prov.: 6. Orthopodomyia bacigalupoi. Chiquitos (?San José de), Agua retirada, Chiquitos Prov.: 16. Haemagogus (S.) lindneri. Lagunillas, Valle Grande Prov.: 6. Psorophora (G.) dimidiata. Naranjo (Los Naranjos): 4. Mansonia(M.) fonsecai; 9. Psorophora (G.) chi- quitana. Pailon (El): 19. Edwardsops boliviensis. Pampa Grande, Florida Prov.: 13. Aedes (H.) aurivittatus. Puerto Suarez, Chiquitos Prov.: 5. Mansonia nitens. CHILE List of Species 1. Anopheles (A.) neghmei Mann, 1950 [ssp. of pseudopunctipennis|. TYPE: Holotype o, Quebrada de Mifiemifie (Tarapaca), elev. 1800 m, Feb-Mar or June 1946, G. Mann F. (Inst. Biol. "Juan Noe,"' Santiago, A-1). BIONOMICS: Larvae in natural and artificial ground waters. 2. Anopheles (A.) noei Mann, 1950 [ssp. of pseudopunctipennis]. TYPE: Holotype o", Oasis of Suca (Tarapaca), elev. 1330 m, Feb-Mar or June 1946, G. Mann F. (Inst. Biol. 'Juan Noe,'' Santiago, A-32). BIONOMICS: Larvae primarily in irrigation waters. 3. Anopheles (Nyssorhynchus) pictipennis Philippi, 1865. TYPE: 20; San- tiago (Santiago), July 1859; Aconcagua province, Sept 1863 (NE). BIONOM- ICS: [Larvae probably in permanent or semipermanent ground waters. | 4. Anopheles (N.) bigotii Theobald, 1901 [=pictipennis]. TYPE: Holotype °, locality not specified, 1894, J.M.F. Bigot (NE). BIONOMICS: [Larvae prob- ably in permanent or semipermanent ground waters. | 5. Anopheles (N.) variegatus (E. Blanchard, 1852); chilensis (R. Blan- oe Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 chard, 1905), nom.nov. [=pictipennis]. TYPE: Lectotype ?, Arquero [Arque- ros?] (MNHP; designation of Belkin, 1968:10). BIONOMICS: [Larvae probably in permanent or semipermanent ground waters. | 6. Psorophora marmorata (Philippi, 1865) [identity doubtful]. TYPE: 2 ¢, 1°, locality not specified (NE). BIONOMICS: [Larvae probably in ground pools. | 7. Aedes (Ochlerotatus) flavipes (Macquart, 1838)[=?albifasciatus]. TYPE: Holotype ?, Concepcion (Concepcion), no other data (MNHP). BIONOMICS: [Lar- vae probably in temporary ground pools or flooded margins of ponds or streams. | 8. Aedes (O.) annuliferus (E. Blanchard, 1852) [=?albifasciatus]. TYPE: Lectotype 2, probably from vicinity of Coquimbo or Illapel (Coquimbo) (MNHP; designation of Belkin, 1968:4). BIONOMICS: [Larvae probably in temporary ground pools or flooded margins of ponds or streams.| _ 9. Aedes (O.) vittatus (Philippi, 1865); philipii Dyar, 1924, new name [= ?albifasciatus]|. TYPE: 2 9, Santiago (Santiago), no other data (NE). BIONOM- ICS: [Larvae probably in temporary ground pools or flooded margins of ponds or streams. | 10. Aedes (O.) colonarius Dyar, 1924 [=?albifasciatus]. TYPE: Holotype ?, Azapa Valley (Tarapaca), June 1912, C.E. Porter (USNM). BIONOMICS: [Larvae probably in temporary ground pools or flooded margins of streams. | 11. Culex (C.) annuliventris (E. Blanchard, 1852) [A distinct species]. TYPE: Lectotype &, possibly from Valdivia (Valdivia) (MNHP; designation of Belkin, 1968:13). BIONOMICS: [Larvae in permanent or semipermanent ground waters. | 12. Culex (C.) apicinus Philippi, 1865. TYPE: Adult, near Santiago (San- tiago), no other data (NE). BIONOMICS: [Larvae probably in permanent or sem- ipermanent ground waters, ponds or streams.| 13. Culex (C.) articularis Philippi, 1865. TYPE: Original material from near Corral (Valdivia) all lost (NE); neotype o’, Casa Pangue (Llanquihue), Dec 1926, R. and E. Shannon (USNM, RB 62 #554; designation of Bram, 1967:30). BIONOMICS: [Larvae in permanent or semipermanent ground waters. | 14. Culex (C.) serotinus Philippi, 1865 [=pipiens complex]. TYPE: , 9, Santiago (Santiago) and Valdivia (Valdivia), no other data (NE). BIONOMICS: [Larvae probably in ground waters contaminated with domestic wastes or in large artificial containers. | 15. Nothodixa chilensis (Alexander, 1913). TYPE: Holotype o, Concepcion (Concepcion), 23 Aug 1904, P. Herbst (NE). BIONOMICS: [Larvae probably in vegetation or flotage on margins of streams, rivulets, seepages and springs. | 16. Nothodixa ensifera (Edwards, 1930). TYPE: Holotype “, Casa Pangue (Llanquihue), elev. about 1000 ft, 4-10 Dec 1926, F.W. Edwards (BM). BIO- NOMICS: [Larvae probably in vegetation or flotage on margins of streams, riv- ulets, seepages and springs. | List of Localities [NOT SPECIFIED] Locality and province not specified, probably near Santiago or Valparaiso: 4. Anopheles (N.) bigotii; 6. Psorophora marmorata. ACONCAGUA [Locality not specified]: 3. Anopheles (N.) pictipennis (also Santiago). Belkin et al: Topotypic South American Mosquitoes 23 CONCEPCION Concepcion: 7. Aedes (O.) flavipes; 15. Nothodixa chilensis. COQUIMBO Arqueros: 5. Anopheles (N.) variegatus. Coquimbo: 8. Aedes (O.) annuliferus (also Illapel). Illapel: 8. Aedes (O.) annuliferus (also Coquimbo). LLANQUIHUE Casa Pangue, frontier station on Chilean side of Rosales Pass, altitude of house about 1000 ft: 13. Culex (C.) articularis; 16. Nothodixa ensifera. SANTIAGO Santiago: 3. Anopheles (N.) pictipennis (also Aconcagua Prov.); 9. Aedes (O.) vittatus; 14. Culex (C.) serotinus (also Valdivia). See also [NOT SPECIFIED]. TARAPACA Azapa Valley: 10. Aedes (O.) colonarius. Mifiemifie, Quebrada de: 1. Anopheles (A.) neghmei. Suca, Oasis of: 2. Anopheles (A.) noei. VALDIVIA Corral: 13. Culex (C.) articularis. Valdivia: 11. Culex (C.) annuliventris; 14. Culex (C.) serotinus (also San- tiago). PARAGUAY _ List of Species 1. Phoniomyia fuscipes (Edwards, 1922). TYPE: Lectotype ?, unspecified locality in Paraguay, probably Asuncion (Asuncion), Fiebrig (BM; designation of Belkin, 1968:24). BIONOMICS: [Larvae probably in leaf axils of bromeliads. ] 2. Psorophora (P.) pallescens Edwards, 1922. TYPE: Lectotype % with attached genitalia mount, unspecified locality in Paraguay, probably Asuncion (Asuncion), Fiebrig (BM: designation of Belkin, 1968:28). BIONOMICS: [Lar- vae probably in temporary ground pools or overflows of ponds and streams.| 3. Psorophora (Janthinosoma) purpurascens Edwards, 1922 [=cyanescens]. TYPE: Holotype, locality and date not specified, probably Asuncion (Asuncion), Fiebrig (HNM). BIONOMICS: [Larvae probably in sunlit temporary ground pools. | 4. Psorophora (J.) fiebrigi Edwards, 1922 [=ferox]. TYPE: Lectotype & with attached genitalia mount, unspecified locality in Paraguay, probably Asun- cion (Asuncion), Fiebrig (BM; designation of Belkin, 1968:26). BIONOMICS: [Larvae probably in temporary ground pools in wooded or shaded areas.]| 5. Psorophora (J.) paraguayensis (Strickland, 1911) [=varipes]. TYPE: Holotype 2, Puerto Max (Boqueron), Jan-Apr 1905, Vezényi (NE). BIONOMICS: [Larvae probably in temporary pools in wooded or shaded areas. | 6. Psorophora (Grabhamia) varinervis Edwards, 1922. TYPE: Holotype 9, locality and date not specified, probably Asuncion (Asuncion), Fiebrig (HNM). 24 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 BIONOMICS: [Larvae probably in temporary ground pools or overflows of ponds and streams.| 7. Aedes (Ochlerotatus) stigmaticus Edwards, 1922. TYPE: Lectotype ¢, Asuncion (Asuncion), 1904, Vezényi (BM; designation of Belkin, 1968:7). BIO- NOMICS: [Larvae probably in temporary ground pools or overflows of ponds and streams.| List of Localities ASUNCION [Locality not specified, probably vicinity of Asuncion]: 1. Phoniomyia fus- cipes; 2. Psorophora (P.) pallescens; 3. Psorophora (J.) purpurascens; 4. Psor- ophora (J.) fiebrigi; 6. Psorophora (G.) varinervis. Asuncion: 7. Aedes (O.) stigmaticus. BOQUERON Puerto Max: 5. Psorophora (J.) paraguayensis. PERU List of Species 1. Anopheles (Stethomyia) acanthotorynus Komp, 1937. TYPE: Holotype & with genitalia slide, Iquitos (Loreto), Apr 1931, R.C. Shannon (USNM, 52020). BIONOMICS: [Larvae probably in deeply shaded stream bed pools.| 2. Anopheles (A.) peruvianus Tamayo, 1907 [=pseudopunctipennis]. TYPE: o', 2, Huacachina (Ica), San Pedro de Lloc (La Libertad), Lima, summer and fall, Chanchamayo (LU). BIONOMICS: [Larvae probably in various small, sun- lit, permanent and semipermanent ground waters with algae.| 3. Trichoprosopon (T.) subsplendens (Martini, 1931) [=digitatum]. TYPE: Holotype 2, Pt. Bermudas [Bermudez], Pichis [Rio] (Pasco), 10/12.03 (NE). BIONOMICS: [Larvae probably in treeholes, cut bamboo or plant parts on the ground. | 4. Trichoprosopon (Runchomyia) hyperleucum (Martini, 1931). TYPE: Ho- lotype 2, Urubamba River (Cuzco) (NE). BIONOMICS: [Larvae probably in tree- holes, cut bamboo, plant parts on the ground, or in the leaf axils and/or flower bracts of Araceae, Marantaceae, Musaceae or bromeliads. | 5. Aedes (Ochlerotatus) epinolus Dyar & Knab, 1914 [=taeniorhynchus]. TYPE: Lectotype ?, Ventanillas (Cajamarca), 4 Feb 1914, C.H.T. Townsend (USNM, 18362; designation of Stone and Knight, 1956a:217). BIONOMICS: [Lar- vae probably in coastal saltmarshes or freshwater pools near the sea.| 6. Haemagogus (Stegoconops) obscurescens Martini, 1931 [=anastasionis]. TYPE: Lectotype 9, Ucayali River (Loreto) (BM; designation of Mattingly, 1955: 28). BIONOMICS: [Larvae probably in treeholes or cut bamboo. | 7. Culex (C.) debilis (Dyar & Knab, 1914) [=apicinus]. TYPE: Lectotype o' with genitalia and 3 legs on slides (597, 598), Matucan [Matucana] (Lima), elev. 7300 ft, June-July 1913, C.T. Brues (USNM, 18361; designation of Stone and Knight, 1957a:47). _BIONOMICS: [Larvae probably in permanent or semi- permanent ground waters. | 8. Culex (C.) escomeli Bréthes, 1920 [=apicinus]. TYPE: o, ?, Arequipa (BA). BIONOMICS: [Probably as for 7. debilis. Belkin et al: Topotypic South American Mosquitoes 20 9. Culex (C.) archegus Dyar, 1929. TYPE: Holotype o, Tarma (Junin), elev. 10,000 ft, 13 July 1928, R.C. Shannon (USNM). BIONOMICS: [Larvae prob- ably in permanent or semipermanent ground waters. | 10. Culex (C.) diplophyllum Dyar, 1929. TYPE: Lectotype o with genitalia slide (2398), Verrugas Canyon (Lima), 5 Apr 1928, R.C. Shannon (USNM; des- ignation of Stone and Knight, 1957a:47). BIONOMICS: [Larvae probably in per- manent or semipermanent ground waters. | 11. Culex (C.) raymondii Tamayo, 1907 [=pipiens quinquefasciatus]. TYPE: o, 2, larva, pupa, Huacachina (Ica) (LU). BIONOMICS: [Larvae probably in ground waters contaminated with domestic wastes or in large artificial contain- ers. | 12. Corethrella inca Lane, 1939 [=ananacola]. TYPE: Holotype ?, Iquitos (Loreto), Mar-Apr 1931, R.C. Shannon (FH). BIONOMICS: [Larvae probably in leaf axils of bromeliads, possibly Ananas magdalenae. | 13. Corethrella maculata Lane, 1939. TYPE: Holotype o, Iquitos (Loreto), Mar-Apr 1931, R.C. Shannon (FH). BIONOMICS: [Larvae probably in ground waters, possibly in leaf axils or treeholes.| 14. Lutzomiops peruviana (Lane, 1939). TYPE: Holotype ?, Iquitos (Lor- eto), Mar-Apr 1931, R.C. Shannon (FH). BIONOMICS: [Larvae probably in swamps. | 7 15. Lutzomiops shannoni (Lane, 1939). TYPE: Holotype o, Iquitos (Lore- to), Mar-Apr 1931, R.C. Shannon (USNM). BIONOMICS: Larvae in a swamp. 16. Dixella andeana (Lane, 1942). TYPE: Holotype o, Iquitos (Loreto), Mar-Apr 1931, R.C. Shannon (USNM). BIONOMICS: [Larvae probably on mar- gins of small streams or permanent or semipermanent ground pools. | 17. Dixella peruviana (Edwards, 1931). TYPE: Holotype “, Verrugas (Lima), 5 May 1928, R.C. Shannon (BM). BIONOMICS: [Larvae probably on margins of small streams or springs. | List of Localities AREQUIPA [Locality not specified]: 8. Culex (C.) escomeli. CAJAMARCA Ventanillas: 5. Aedes (O.) epinolus. CUZCO Urubamba (Rio): 4. Trichoprosopon (R.) hyperleucum. ICA Huacachina: 2. Anopheles (A.) peruvianus (also San Pedro de Lloc (La Li- bertad) and Lima); 11. Culex (C.) raymondii. J UNIN Tarma: 9. Culex (C.) archegus. LA LIBERTAD San Pedro de Lloc: 2. Anopheles (A.) peruvianus (also Huacachina (Ica) and Lima). 26 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 LIMA [Locality not specified, probably vicinity of Lima]: 2. Anopheles (A.) peru- vians (also Huacachina (Ica) and San Pedro de Lloc, La Libertad). Matucuna: 7. Culex (C.) debilis. Verrugas: 10. Culex (C.) diplophyllum (Verrugas Canyon); 17. Dixella pe- ruviana. LORETO Iquitos: 1. Anopheles (S.) acanthotorynus; 12. Corethrella inca; 13. Core- thrella maculata; 14. Lutzomiops peruviana; 15. Lutzomiops shannoni; 16. Dix- ella andeana. Ucayali (Rio): 6. Haemagogus (S.) obscurescens. PASCO ! Puerto Bermudez, Rio Pichis: 3. Trichoprosopon (T.) subsplendens. URUGUAY List of Species 1. Psorophora (Grabhamia) leucocnemis Martini, 1931 [?=varinervis]. TYPE: Holotype 9, Montevideo (Montevideo), 19 Mar 1927, Vogelsang coll. 8846 (BM). BIONOMICS: [Larvae probably in temporary ground pools or overflows of ponds or streams. | REFERENCES CITED Belkin, John N. 1968. Mosquito Studies (Diptera, Culicidae). IX. The tbe specimens of New World mosquitoes in European museums. Amer. Entomol. Inst., Contrib. 3(4). 69 p. Belkin, John N., R. X. Schick and $.J. Heinemann 1965. Mosquito Studies (Diptera, Culicidae). V. Mosquitoes originally de- scribed from Middle America. Amer. Entomol. Inst., Contrib. 1(5): 1-99, 1966. Mosquito Studies (Diptera, Culicidae). VI. Mosquitoes originally de- scribed from North America. Amer. Entomol. Inst., Contrib. 1(6): 1-39. Bram, Ralph A. 1967. Classification of Culex subgenus Culex in the New World. U.S. Nat. Mus., Proc. 120:1-122. Castro, Manuel and M. Bressanello 1952. Revision de las especies de Taeniorhynchus (Rhyncotaenia) (Dipt. Cul.). Rev. Brasil. Biol., Rio de Janeiro 12:229-246. Garcia, Miguel and R.A. Ronderos (1962) 1963. Sobre la validez de Aedes (Ochlerotatus) patersoni Shannon y del Ponte, 1927 (Diptera-Culicidae). Soc. Entomol. Argent., Rev., Buenos Aires 25:29-33. Peters, T. Michael 1968. Mosquito Studies (Diptera, Culicidae). X. Dixinae originally described Belkin et al: Topotypic South American Mosquitoes at from North America. Amer. Entomol. Inst., Contrib. 4(1):1-7. Stone, Alan and K. L. Knight : 1955. Type specimens of mosquitoes in the United States National Museum: I. The genera Armigeres, Psorophora and Haemagogus (Diptera, Cu- licidae). Wash. Acad. Sci., J. 45:282-289. 1956. Type specimens of mosquitoes in the United States National Museum: III. The genera Anopheles and Chagasia (Diptera, Culicidae). Wash. Acad. Sci., J. 46:276-280. 1957a. Type specimens of mosquitoes in the United States National Muse- um: IV. The genus Culex (Diptera, Culicidae). Wash. Acad. Sci., J. 47:117-126. 1957b. Type specimens of mosquitoes in the United States National Muse- um: V. The Sabethini (Diptera, Culicidae). Wash. Acad. Sci., J. 47: 117-120. | 1957c. Type specimens of mosquitoes in the United States National Muse- um: VI, Miscellaneous genera, addenda, and summary. Wash. Acad. Sci., J. 47:196-202. | Umana, A.C., R.L. Heredia and J.C. Siquot 1959. Estudios sobre el Anopheles (N.) albitarsis en la Argentina (Nota previa). Primeras Jornadas Entomoepidemiologicas Argentinas, Buenos Aires, part 2, p. 609-618. CLASSIFIED INDEX AEDEOMYIA [1] stigmaticus, 24 *squamipennis, 12 AEDES (HOWARDINA) [3] aurivittatus, 20 delpontei, 20 vanemdeni, 20 AEDES (OCHLEROTATUS) [16] albifasciatus annuliferus, 22 colonarius, 22 flavipes, 22 #philipii, 22 vittatus, 22 angustivittatus meprai, 13 bejaranoi, 20 crinifer iguazu, 14 lynchii, 13 tapinops, 13 milleri araozi, 14 oroecetor, 20 patersoni, 14 raymondi, 14 scapularis *confirmatus, 14 taeniorhynchus epinolus, 24 ANOPHELES (ANOPHELES) [8] annulipalpis, 9 fluminensis holmbergi, 9 pseudopunctipennis *argentinus, 9 neghmei, 21 noei, 21 patersoni, 10 peruvianus, 24 tucumanus, 10 ANOPHELES (KERTESZIA) [1] boliviensis, 19 ANOPHELES (NYSSORHYNCHUS) [10] *albitarsis, 10 argyritarsis rooti, 10 evansae, 10 noroestensis clarki, 10 pictipennis, 21 bigotii, 21 #chilensis, 21 variegatus, 21 28 Contrib. Amer. Ent. Inst., vol. 4, no. 1, 1968 triannulatus *bachmanni, 10 davisi, 10 perezi, 10 ANOPHELES (STETHOMYIA) [1] acanthotorynus, 24 bejaranoi, 16 intrincatus, 16 martinezi, 16 misionensis, 16 orfilai, 16 pavlovskyi, 16 COQUILLETTIDIA (RHYNCHOTAENIA)| CULEX (subgenus unknown) [1] [3] fasciolata, 12 nitens, 19 venezuelensis araozi, 12 CORETHRELLA [5] ananacola inca, 25 appendiculata arborealis, 16 maculata, 25 puella, 16 quadrivittata, 16 CULEX (ALLIMANTA) [1] *tramazayguesi, 15 CULEX (CULEX) [23] ameliae, 14 annuliventris, 22 apicinus, 22 debilis, 24 escomeli, 24 archegus, 20 articularis, 22 bidens, 20 interfor, 14 brethesi, 15 castroi, 15 diplophyllum, 25 *dolosus, 15 bonariensis, 15 eduardoi, 15 fernandezi, 15 hepperi, 15 lahillei, 15 maxi, 15 pipiens autumnalis, 15 raymondii, 25 serotinus, 22 saltanensis, 15 CULEX (LUTZIA) [1] bigoti patersoni, 14 CULEX (MELANOCONION) [7] aliciae, 16 oblita, 16 DIXELLA [3] andeana, 25 argentina, 17 peruviana, 25 EDWARDSOPS [1] boliviensis, 20 HAEMAGOGUS (STEGOCONOPS) [5] anastasionis obscurescens, 24 capricornii petrocchiae, 14 spegazzinii, 14 lindneri, 20 uriartei, 14 LIMATUS [1] durhamii exhibitor, 11 LUTZOMIOPS [3] davisi, 17 peruviana, 25 shannoni, 25 /MANSONIA (MANSONIA) [1] indubitans fonsecai, 19 NOTHODIXA [4] atrovittata, 17 chilensis, 22 ensifera, 22 nitida, 17 ORTHOPODOMYIA |[1} fascipes bacigalupoi, 20 PHONIOMYIA [2] fuscipes, 23 muehlensi, 11 i PSOROPHORA (GRABHAMIA) [6] *confinnis, 13 chiquitana, 20 dimidiata, 20 paulli, 13 varinervis, 23 leucocnemis, 26 PSOROPHORA (JANTHINOSOMA) [8] circumflava, 20 Belkin et al: Topotypic South American Mosquitoes cyanescens purpurascens, 23 ferox centrale, 13 fiebrigi, 23 lutzii chaquensis, 13 melanota, 20 — varipes bruchi, 13. paraguayensis, 23 PSOROPHORA (PSOROPHORA) [5] ciliata lynchi, 12 holmbergii, 13 agoggylia, 13 pallescens, 23 stigmatephora, 13 PHOROPHORA (subgenus unknown) [1] marmorata, 22 SABETHES (SABETHES) [1] albiprivus neivai, 12 SAYOMYIA [2] australis, 16 elnorae, 16 TOXORHYNCHITES (ANKYLORHYN- CHUS) [1] hexacis, 19 TOXORHYNCHITES (LYNCHIELLA) [5] cavalierii, 10 guadeloupensis arborealis, 10 tucumanus, 10 29 haemorrhoidalis lynchi, 11 separatus, 11 TRICHOPROSOPON (CTENOGOELDIA) 1] magnum, 19 TRIC HOPROSOPON (RUNCHOMYIA) [2] hyperleucum, 24 paranensis, 11 TRICHOPROSOPON (TRICHOPROSO- PON) [1] digitatum subsplendens, 24 URANOTAENIA [77] capitis, 12 lanei, 12 monilis, 12 *nataliae, 12 *pulcherrima, 12 elnora, 12 urania, 12 WYEOMYIA (DAVISMYIA) [2] *petrocchiae, 11 monoleua 11 WYEOMYIA (DENDROMYIA) [2] belkini, 11 : melanocephala typharum, I1 WYEOMYIA (MENOLEPIS) [1] leucostigma leontiniae, 11 WYEOMYIA (NUNEZIA) [1] lateralis, 11 eat o “*< in | im . é wits iM So hie | d int ars id nine “ee Ditty’. P. Mala HDS VtY ah of Perit 4 piles ay yep ein a ‘ ae | ages nai Contributions of the American Entomological Institute Volume 4, Number 2, 1969 MOSQUITO STUDIES (Diptera, Culicidae) XII. A revision of the Neotropical subgenus Howardina of Aedes. By O. G. W. Berlin a NSO Wi qas AUG 8 14969 C/BRARIED CONTRIBUTIONS of the AMERICAN ENTOMOLOGICAL INSTITUTE The Contributions are for larger papers on insects. Each paper is a separate number, with separate pagination and index. Separate numbers aggregating about 500 pages constitute a volume. Issues appear irregularly, as suitable manuscripts are available. Copies are sold separately or in subscriptions to complete volumes. Complete volumes are $12.00. The price of separate numbers varies. Subscribers are billed for each volume with its beginning number, and receive the parts as issued. Orders for separate numbers that total less than $8.00 must be accompanied by payment. 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CONTENTS INTRODUCTION. . fs MATERIAL AND METHODS . ) A GENERAL CONSIDERATIONS . 5 Taxonomic characters . 5 Bionomics 6 Distribution . 7 Systematics and Evolution , g Affinities . 7 Medical Importance . 8 TAXONOMIC TREATMENT . 8 Subgenus Howardina sheca HA tengo Sh GP eB RR died nik gibt ag ee Partie re 1 Nh A ae ian el Ur, i er ie Rae he ate bik baat tte ag iggy abe bare PRY OTR CEO 2 38 nse os), RN Oa a hei wa es eee Keys to species. . Cede Wai i ET isu omen tae RE ee Cine a ue ae 1. Aedes (H.) fulvithorax (Lutz) .. mae Be easier OAM ee cmon CPN 2. Aedes (H.) septemstriatus Dyar and Knab pa Alias ae RAISE Aa Ree eR 3. Aedes (H.) arborealis Bonne-Wepster and Bonne. . ....... 22 OR ORUTE ) CUPS GOGO DBD oo ee ON re ae a 5 eres. ) 5p. SOS POP oF. He erie ae ie eae 1Contribution from project “Mosquitoes of Middle America” supported by U.S. Public Health Service Research Grant AI-04379 and U.S. Army Medical Research and Development Command Research Contract DA-49-193-MD-2478. Based on Ph.D. dissertation submitted to the University of California, Los Angeles and supported in part by National Science Foundation Grant GB 3871 for field studies. Department of Zoology, University of California, Los Angeles, California 90024. Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Walkeri Section Keys to groups and soecies Walkeri Group. . . 6. Aedes (H.) walkeri (Theobald) 7. Aedes (H.) argyrites Dyar and Nunez Tovar ; Aurites Group . . poe eae 8. Aedes (H.) aerihes (Theobald). 9. Aedes (H.) grabhami,n.sp. . 10. Aedes (H.) inaequalis (Grabham) 11. Aedes (H.) stenei Thompson . Ioliota Group . . 12. Aedes (H.) joliota Dyvar and oe 13. Aedes (H.) cozumelensis Diaz Najera . Albonotatus Group. . . 14. Aedes (H.) albonotatus (Coquille) 15. Aedes (H.) bahamensis, n.sp. Busckii Group . . 16. Aedes (H.) busckti (Coquillett) Sexlineatus Section . Keys to groups and species Quadrivittatus Group . . 17. Aedes (H.) quadrivittatus (Coquillett) 18. Aedes (H.) lorraineae, n.sp. 19. Aedes (H.) brevis, n.sp. . 20. Aedes (H.) spinosus,n.sp. . Sexlineatus Group . . 21. Aedes (H.) sexlineatus (Theobald). Whitmorei Group. . . 22. Aedes (H.) whitmorei Dunn . 23. Aedes (H.) pseudodominicii Komp. Allotecnon Group. 24. Aedes (H.) allotecnon Kumm, Komp and Ruiz . 25. Aedes (H.) guerrero, n.sp. 26. Aedes (H.) guatemala, n.sp. Eleanorae Group . . 27. Aedes (EH) eleanorae, n. sp. 28. Aedes (H.) marinkellei, n.sp. . 29. Aedes (H.) leei, n.sp. . 30. Aedes (H.) osornoi, n.sp. 31. Aedes (H.) ecuadoriensis, n.sp. . 32. Aedes (H.) brevivittatus, n.sp. Aurivittatus Group . . 33. Aedes (H.) aurivittatus Cerqueira ; 34. Aedes (H.) martinezi,n.sp.. 35. Aedes (H.) vanemdeni Martini REFERENCES CITED FIGURES INDEX TO SCIENTIFIC NAMES Berlin: Subgenus Howardina 3 INTRODUCTION The exclusively Neotropical subgenus Howardina has been recognized as a distinct entity in the widespread genus Aedes since Dyar’s classification of the American spe- cies of the genus (Dyar 1920:103-106) but no attempt has been made to date to de- velop an internal classification of this subgenus. Nearly all species assigned to How- ardina have been characterized entirely on the pattern of ornamentation of the adults. The immature stages were largely unknown. In recent years, considerable as- sociated reared material has accumulated and it is now possible to attempt a compre- hensive revision of the subgenus based on all stages. The taxonomic history of Howardina is brief and relatively simple except for con- fusion with other generic group taxa (Finlaya, Haemagogus, Stegomyia, etc.) which need not be detailed here. Howardina was proposed by Theobald (1903:287) as a distinct genus with 2 included species, walkeri and greenii, and with the following definition: ‘“‘Head clothed with flat and narrow curved scales, the latter forming a median area as in Aedes. Scutellum with narrow curved scales; the midlobe with four posterior border-bristles. Wings with lateral vein-scales large for the size of the wings, long, rather thin, the median vein-scales small. Palpi of the 2 minute, four-jointed; the penultimate joint long, longer than the two basal ones; apical joint very minute. 6 unknown. All small species, and with ornamented thorax.” Dyar (1905:44) selected walkeri as the type species of Howardina. Coquillett (1906:312) recognized the im- portance of the simple claws, a character mentioned in the original description of walkeri. By 1910 Theobald recognized 7 species in the genus, described by himself and by Grabham, but 3 of these were subsequently removed to the genus (later sub- genus) Finlaya. Howard, Dyar and Knab (1912a:65,70) synonymized Howardina with Aedes but Dyar (1920:103) elevated it to subgeneric rank within Aedes on the basis of male genitalic characters (rudimentary claspette). All other contributions be- tween 1910 and 1932 were merely descriptions of individual species. Edwards (1932: 155-156) in his classic revision of the mosquitoes of the world listed 13 species in the subgenus Howardina and gave the following brief diagnosis which has been followed by all authors to the present: “‘most closely related to Finlaya, from which it differs chiefly in the simple claws of the ¢; less prominent eighth sternite of 9 abdomen; and less developed claspette of 6 hypopygium, the stem being very short and the append- age bristle-like. Vertex in all species with a median stripe of narrow scales reaching forward to eyes; scutellar scales narrow. Thorax conspicuously ornamented with white or golden lines. Larva.- Similar to that of Finlaya; a more or less distinct small chitinous plate present on each side of the ventral brush.’ Subsequent to Edwards’ revision more confusion than progress has been accomplished, particularly in errone- ous identifications of species. In the world catalog of mosquitoes (Stone, Knight and Starcke 1959:175-176) 16 species were recognized on the basis of published records. Included in Edwards’ (1932:155-156) list of species of Howardina were dominicii and whitmorei for which Komp (1936:71) subsequently erected the subgenus So- peria, with dominicii as the type species. Unfortunately Komp misidentified domi- nicii which in reality is conspecific with sexlineatus in the subgenus Howardina. Komp based the new subgenus primarily on the complete absence of a claspette in the male genitalia. Soperia has been considered as a distinct subgenus to the present and contained 3 species in the world catalog (Stone, Knight and Starcke 1959:173), one of which, tracei, was synonymized with busckii in the subgenus Howardina by Belkin, Schick and Heinemann (1965:33). 4 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 In the present study only the subgenus Howardina is recognized; the species for- merly placed in Soperia of authors form a distinct group included in the Sexlineatus © Section, 1 of 3 primary divisions of Howardina. A total of 35 species is now recog- nized in the subgenus: 5 species in 1 group in the Fulvithorax Section; 11 species in 5 groups in the Walkeri Section; and 19 species in 6 groups in the Sexlineatus Sec- tion. I wish to express my gratitude to Dr. John N. Belkin for suggesting this project and directing the study. I am indebted to Dr. Alan Stone of the U.S. National Muse- um for the loan of material and for information on type specimens; to Dr. P.F. Mat- tingly for the loan from the British Museum; and to Dr. A. Martinez, Dr. A. Spielman and Dr. A. Diaz Najera for specimens from Argentina, Bahamas and Mexico respec- tively. I also thank Pedro Galindo, Gorgas Memorial Laboratory, Panama, and Dr. I. Goodbody, Department of Zoology, University of West Indies, Kingston, Jamaica, for the courtesies extended to me during field trips to these countries. Finally I wish to thank Dr. R.X. Schick, Dr. S. Sirivanakarn, Dr. T.J. Zavortink and Miss Sandra Heinemann for the valuable suggestions and assistance and to Miss Sheila Bernstein for the preparation of the final copy for reproduction. MATERIAL AND METHODS MATERIAL. This study is based largely on extensive collections of adults and as- sociated immature stages accumulated at the University of California, Los Angeles for the project ‘““Mosquitoes of Middle America’’ (Belkin, Schick et al, 1965) and also on material from the U.S. National Museum and the British Museum (Nat. Hist.) where most of the types of previously described species are deposited. Additional critical material was obtained specifically for this study in Jamaica in November and December 1966 following the methods of collection and rearing outlined by Belkin, Hogue et al (1965). For this revision a total of 10,949 specimens were studied, including 1,240 6, (3,527 2, 2,412 pupae, 3,770 larvae, with 1,796 individual rearings (810 pupal, 645 larval, 341 incomplete). TAXONOMIC PROCEDURE. The taxonomic methods used in this study are the Classical phylogenetic ones, simple in principle but difficult in practice. Following the recognition of species on the basis of a series of constant correlated characters in all stages, the different species were compared minutely and those showing the greatest © similarities in all stages were interpreted as members of primary phyletic lines or spe- cies groups, called simply Groups here. These groups were compared in turn to estab- lish the more inclusive Sections within the subgenus. DESCRIPTIONS. The method of presentation, terminology and abbreviations used in the descriptions of taxa in general follow Belkin (1962). A few special terms used for the pattern of ornamentation of adults are explained in the section on taxo- nomic characters in the following chapter. ; ILLUSTRATIONS. All available stages of every species are figured, the larvae and pupae in full and the adults in pertinent details only. Each illustration is based on a topotypic specimen, whenever available, but the character states shown represent the modal condition determined usually from at least 10 topotypic specimens. In the il- lustrations of the ornamentation of mesonotum and pleuron, dotted lines represent golden or creamy scales and dashed lines denote white or silvery scales. | Berlin: Subgenus Howardina 5 DISTRIBUTION DATA. In the distribution lists, the arrangement of countries is from north to south and west to east and the localities in each country are listed al- phabetically under the major political subdivisions. Included in these lists are all the records of material | have examined as well as published records from the literature which I consider reliable. In these lists, P and L denote whole pupa and larva and p and |, pupal and larval skins respectively. DEPOSITORIES. The abbreviations used by Stone, Knight and Starcke (1959) for institutions in which types are deposited are followed here to indicate the loca- tion of the material studied. To these are added the Institute of Jamaica [IJ] and the Department of Zoology, Los Angeles, California [UCLA]. The bulk of the material will remain at UCLA. The types of all new species described here will be deposited in the U.S. National Museum. GENERAL CONSIDERATIONS TAXONOMIC CHARACTERS. The following discussion is limited to external morphological characters. Other important taxonomic characters, ecological and geo- graphical, are discussed below in the sections on Bionomics and Distribution respec- tively. : hae, In the adults, the most important taxonomic characters pertain to the pattern of ornamentation of the mesonotum. As indicated in the diagnosis of the subgenus, the _ pattern usually consists of lines of light scales superimposed’ on a background of dark scales. Each section, each group and most species have a distinctive pattern of orna-: mentation. It was necessary to develop a special terminology for these light lines as no suitable one was available. This terminology is illustrated in figs.8 and 9 and ex- plained below. The acrostichal line is the single median longitudinal mesonotal line and is developed only in some members of the Fulvithorax Section; it ends in front of the prescutellar space and is more or less distinctly continued caudad as a pair of lateral prescutellar lines. The median prescutellar line is developed only in the Walkeri Section and is in the median longitudinal position in the “‘bare’’ prescutel- lar space. The inner dorsocentral lines are located mesad of the dorsocentral row of bristles in a submedian longitudinal position on each side; these lines reach the scu- tellum only in the Sexlineatus Section; the posterior parts (prescutellar) of these lines more or less correspond to the lateral prescutellar lines found in some members of the Fulvithorax Section. The outer dorsocentral lines are located laterad of the dorsocentral row of bristles in a sublateral longitudinal: position; they are repre- sented to a varied degree in all sections and usually reach the lateral lobes of the scu-: tellum in the Walkeri and Sexlineatus Sections. The lateral marginal lines border the mesonotum on each side to a varied degree from the anterior promontory to the wing root; they may be broken into one or more prescutal and antealar seg- ments. Transverse diagonal posterior fossal lines may connect the lateral marginal to the outer dorsocentral on each side. Longitudinal supraalar lines may also be devel- oped and connected to the lateral marginal lines on each side caudad of the scutal an- gle. In fulvithorax and to a lesser extent in arborealis of the Fulvithorax Section, the ornamentation is primarily light and may be considered to have been developed by the expansion and fusion of the light lines which are discrete in the other sec- tions, particularly in the Walkeri Section. Other significant taxonomic features of adult ornamentation are: (1) on the head, scaling of the vertex and of the torus and first flagellar segment of the antenna; (2) 6 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 scaling of the paratergite; (3) distribution of the scales on the pleuron, particularly ppn, stp and mep; (4) on the legs, light scaling of the foretibia, midfemur and hind- femur, and the distribution and extent of light scales on the tarsal segments. The comparatively simple male genitalia of Howardina snow relatively few obvious taxonomic characters; even specific differences are usually subtle and sometimes not apparent. The reduced claspette is more or less characteristically developed in each section and is absent in the Whitmorei Group. Other significant taxonomic characters appear to be: (1) extent of separation of the base of the sidepieces midsternally, (2) width and length of the aedeagus in relation to the proctiger and (3) the shape of the aedeagus in dorsal aspect. Only 3 species (bahamensis, whitmorei and pseudodomi- nicii) have genitalia strikingly differentiated from the others in features of the side- pieces. The pupae are quite similar in the subgenus but a few characters separating the 3 sections have been noted in: (1) development of cephalothoracic hair 5-C, (2) posi- tion of abdominal hair 2-[V-VI and (3) development of abdominal hair 9-VI. On the other hand, reliable group characters within a section have not been found and spe- cific differences are rather subtle and not always reliable. The 4th-instar larvae show many taxonomic characters at the section and group levels usually concordant with those of adult ornamentation, and many reliable spe- cific differences. Among the important taxonomic characters are: (1) presence or ab- sence of marginal spicules on the anal saddle, (2) comb scales in a single row or in a patch, (3) ventral brush with 5 or 6 pairs of hairs, (4) prothoracic hair 13-P present or absent, (5) presence or absence of integumentary spicules on the thorax and/or abdomen, and (6) numerous features in the development of hairs, particularly on the head. BIONOMICS. All members of the subgenus, as far as known, are container breed- ers. Immature stages have been reported from rockholes, treeholes, broken bamboo, plant parts on the ground, leaf axils of bromeliads and aroids, and flower bracts of heliconias. A few species invade artificial containers of various types. Only the Walkeri Section utilizes the full range of breeding habitats; the Fulvithorax Section is apparently restricted to treeholes and broken bamboo and the Sexlineatus Sec- tion is known to breed only in bromeliad axils. : In common with other ornamented mosquitoes, species of Howardina are appar- ently largely diurnal in activity (Bates 1944:159-170; 1949:16,61; Carpenter, Galin- do and Trapido 1952:163; Trapido, Galindo and Carpenter 1955:530,537,538; For- attini 1965:401-403). To supplement the published information, I made a full 24- hour observation on the biting activity of females near Porus, Manchester, Jamaica at an elevation of approximately 1000 ft, where 4 species of Howardina were pres- ent. Biting-landing collections were made at hourly intervals at ground level under bromeliad laden trees. As seen in fig.7 there appear to be specific or group differ- ences in activity since walkeri shows 2 definite peaks, a higher one in the morning, and members of the Aurites Group show only a moderate afternoon peak. Although a number of species are known to bite man in forested areas, the definite altitudinal stratification of different species suggests that their natural hosts are a variety of vertebrates characteristic of a particular stratum in the forest. Members of the Quadrivittatus Group have been collected mainly in the forest canopy (Trapido and Galindo 1957:122,123,124,125,131), while members of the Fulvithorax Group are predominantly forest floor biters (Trapido and Galindo 1957:131); members of the Sexlineatus Group (Trapido and Galindo 1957:132) and Whitmorei Group (Bates 1944:166), on the other hand, demonstrate inconsistent behavior. Berlin: Subgenus Howardina 7 DISTRIBUTION (fig.1). Howardina is strictly a Neotropical taxon, centered in the American Mediterranean region in the sense of Belkin (1962:562). At present its known northern limit is Grand Bahama Island in the east and the Sierra Madre del Sur and the southern end of the Sierra Madre Oriental in Mexico in the west. The southern limit appears to be in northern Argentina along the Andes; the subgenus ap- parently does not reach the mouth of the La Plata river in the east. The distributions of the 3 sections are largely complementary. The Walkeri Sec- tion is almost entirely confined to the islands of the Caribbean area with only a slight overlap with the other 2 in northern Venezuela and Trinidad. The Fulvithorax Section is found primarily in eastern South America but with a significant overlap with the Sexlineatus Section in southern Central America. The Sexlineatus Section is primarily associated with the mountain regions of Central America and western South America. SYSTEMATICS AND EVOLUTION. The 35 species recognized in the present study fall into 12 largely well-defined, discrete groups of 1 or more species each. On the basis of morphological similarity the 12 groups form 3 very distinct sections in the subgenus, the Fulvithorax Section with only 1 group (Fulvithorax) of 5 species (1 not named), the Walkeri Section with 5 groups (Walkeri, Aurites, Ioliota, Albo- notatus, and Busckii) of 11 species, and the Sexlineatus Section with 6 groups (Quadrivittatus, Sexlineatus, Whitmorei, Allotecnon, Eleanorae and Aurivittatus) of 19 species. As indicated in the preceding discussions, as well as in the taxonomic treatment of each section, there is practically no overlap in diagnostic morphological characters among the 3 sections, little overlap in distribution, and strong differentiation in the breeding sites between Fulvithorax Section and the Sexlineatus Section. It appears, therefore, that the 3 sections represent 3 old phyletic lines. In spite of this clear-cut divergence, there is a remarkable basic similarity in all stages in all 3 sections. This is undoubtedly indicative of true affinity and some kind of monophyletic origin. Two general explanations for the development of 3 major phyletic lines in Howardina are possible. First, would be serial or independent devel- opment of 2 of the sections from the third; this is not plausible because each of the lines has retained primitive features absent in 1 or 2 of the others. Second, would be a tripartite splitting of one ancestral species. This interpretation is supported by the present distributions of the 3 sections and I therefore suggest on this basis that the original stock was divided into 3 segments: (1) one in the Caribbean, giving rise to the Walkeri Section, (2) one in Central America, giving rise to the Sexlineatus Sec- tion, and (3) one in northeastern South America, giving rise to the Fulvithorax Sec- tion. Subsequent speciation and evolution have followed different patterns in each sec- tion, probably because of different environmental conditions. In the Fulvithorax Section, which occupies primarily eastern South America where relatively few bar- riers to dispersal have existed, at least in relatively recent geological times, there has been little differentiation and speciation. The Walkeri Section shows typical complex insular speciation of a variety of divergent types, including relict forms. In the rather homogeneous Sexlineatus Section the extensive speciation reflects the barriers and environmental diversity associated with the complex orogenies of the cordilleras of western America. AFFINITIES. The general similarity between the subgenera Howardina and Fin- laya has been recognized by many culicidologists, notably Edwards (1932:156), and as indicated in the introduction, some species of Finlaya were originally included in Howardina, primarily on the basis of the ornamentation of the adults. The chief dif- 8 Contrib. Ate, But Inch aca danas 2, 1969 ferences from Finlaya are the untoothed claws and less prominent sternite VIII in the females and the claspette poorly developed or completely absent in the males. In the last feature, Howardina shows little difference from the Oriental dissimilis sub- group of Finlaya (Knight and Marks 1952:516-517). The immature stages of How- ardina also bear a general resemblance to those of Finlaya and are very similar to those of some species of the genus Haemagogus, which is also restricted to the Neo- tropical region and may have evolved from a Finlaya stock “‘through some such spe- cies as A. (F.) leucocelaenus”’ (Edwards 1932:178). The following unusual combina- tion of features in the larvae of Howardina may be diagnostic for the subgenus: ab- dominal hair 12-I never developed, hair 9-I always ventrad of hair 7-I, and tubercles of prothoracic hairs 5-7-P usually distinctly joined (except in Fulvithorax Section). Little can be done now to clarify the affinities of Howardina chiefly because the immature stages of the majority of annectent groups in the tribe Aedini are very poorly known both in the Old World and the New World. In spite of some apparent slight overlap with the subgenus Finlaya and the genus Haemagogus, Howardina is a very distinct group, and should be retained as a subgenus of Aedes. MEDICAL IMPORTANCE. Little information is available on the medical impor- tance of species of Howardina. A few of the more abundant species were investigated in connection with studies on the vectors of jungle yellow fever. Experimental stud- ies with fulvithorax showed that while it could be infected with and maintain the virus, it was not able to transmit it naturally through bites (Davis and Shannon 1931; Whitman and Antunes 1937). Similar results were obtained with other species of Howardina by Boshell-Manrique and Osorno-Mesa (1944), Bugher, Boshell-Manrique et al (1944) and Aitken (1960). Bates (1949:205) has suggested that the incubation period of this virus in some mosquitoes may be a long one and it is possible that Howardina species may transmit the yellow fever virus if the incubation period is prolonged at favorable temperatures. Galindo (1964) reported the isolation of an unidentified virus from guadrivittatus in Panama. Since some species of Howardina, especially members of the Walkeri Section (Walkeri, Aurites and Busckii Groups), attack man readily and are very abundant, a more thorough investigation of the role of these species as vectors of arboviruses should be made. TAXONOMIC TREATMENT Subgenus HOWARDINA Theobald 1903.Howardina Theobald, 1903:287. TYPE SPECIES: Culex walkeri Theobald, 1901, Jamaica; the second of 2 included species, selection of Dyar (1905a:49). 7 1936.Soperia Komp, 1936b:71. TYPE SPECIES: Aedes dominicii Rangel and Romero-Sierra, 1907, Venezuela, original designation. NEW SYNONYMY. Aedes (Howardina) of Dyar (1920:10); Bonne and Bonne-Wepster (1925 :366); Dyar (1928:103); Shannon (1931:125); Edwards (1932:155); Gerry (1932:40); Lane (1939:98); Anduze (1941: 14); Bates (1949:320); Lane (1949:255,261); Del Ponte, Castro and Garcia (1951:231,239); Lane (1953:706); Levi-Castillo (1953:40); Horsfall (1955:471); Martinez and Prosen (1955:21- 22); Peyton, Galindo and Blanton (1958:95-100); Stone, Knight and Starcke (1959:173-174); Martinez, Prosen and Carcavallo (1960:25); Belkin (1962:340); Stone (1963:127); Prosen, Car- cavallo and Martinez (1964:103); Forattini (1965:397-401); Cova Garcia, Sutil and Rausseo (1966a:in part; 1966b:333); Porter (1967:38); Stone (1967:208). Berlin: Subgenus Howardina 9 Aedes (Soperia) of Lane (1939:98); Anduze (1941:14); Senevet and Quievreux (1941:258); Andu- ze (1943:193; 1947:357); Lane (1953:700); Levi-Castillo (1953:40); Komp (1956:38); Stone, Knight and Starcke (1959:173); Forattini (1965 :404,460,462); Cova Garcia, Sutil and Rausseo (1966b:338-339). , Howardina of Blanchard (1905:399,415); Dyar (1905:49); Theobald (1905:21); Theobald and Grabham (1905:7), Coquillett (1906b:16,26); Dyar and Knab (1906b:188); Theobald (1907: 21,149,214); Coquillett (1910:553); Theobald (1910:220); Surcouf and Gonzalez-Rincones (1911:102,139-141; 1912:276). Aedes in part of Howard, Dyar and Knab (1912:65,70; 1917:613); Dyar (1918:71). FEMALES (fig.16). Small to medium sized ornate species; usually blackish to dark brown with white, silver, creamy, yellowish or golden markings on head, thorax and abdomen and with white basal tarsal markings. Head: Eyes distinctly separated be- tween antennae. Decumbent scales predominantly broad and flat; narrow scales usu- ally largely restricted to median longitudinal light stripe of varied width running from lower end of interocular space to occiput; lower end of interocular space always with a tuft of very long outstanding narrow bristlelike striated scales; orbital line of nar- row light scales, varied in width and length, frequently developed. Erect scales short, usually forked apically, restricted to occiput or extending on to the disc and sides of vertex. One pair (rarely 2) of interorbital bristles always differentiated; a single row of orbitals, upper usually 4(3-6) pairs, heavier, longer and more closely spaced than lower. Clypeus bare, deep brown to black. Proboscis slender; subequal to or distinct- ly longer than forefemur; entirely dark scaled; with a few basal bristles. Palpus al- ways short, about 0.15 to 0.2 of proboscis; 5-segmented, segments 1 and 2 anky- lozed, segment 5 minute; segments 1 and 5 without scales, remainder entirely or pre- dominantly dark scaled, light scales usually restricted to apex of segment 4. Antenna distinctly shorter than or subequal to proboscis; torus dark brown to black, with light to dark scales always present, sometimes indistinct, usually restricted to mesal surface; flagellar segment 1 only slightly longer than segment 2, always with a few small scales on mesal surface; flagellar segments 2-13 usually with 6 moderate bris- tles in basal whorls. Thorax: Integument dark brown to almost black. Mesonotum usually with a background of short, narrow curved scales auburn to black in color and with a varied pattern of lines of light scales (except in fulvithorax and arborealis in which light scales predominate) as follows: (1) median acrostichal, (2) inner dorso- central, (3) outer dorsocentral, (4) lateral marginal, (5) prescutal, (6) posterior fossal, (7) supraalar, (8) median prescutellar and (9) lateral prescutellar. Acrostichal bristles usually not developed on disc (present in busckii); dorsocentrals, prescutellars and supraalars always present, variously developed; 1 posterior fossal and 1 parascutellar always developed. Median scutellar lobe with 4,5 large marginal bristles and with dark broad flat scales laterally and narrow light scales in varied median longitudinal line; lateral lobe with 3-5 large marginal bristles and usually with narrow scales only. Para- tergite bare or with scales. Bristles present on apn, ppn, ppl, psp, stp, pra and upper mep; lower mep bristles always absent; lowermost stp bristles distinctly ventrad of lower margin of mep. Pleural scaling dingy white, white or silvery; always present on apn, ppl, base of pra, stp and mep; ppn, psp, and ssp with or without scales. Legs: Coxae with light scales on external surface; trochanters with mesoventral light scales. . White or silvery knee spots prominent on midleg and hindleg, slightly indicated on foreleg. Midfemur and hindfemur with or without distal light spot on anterior surface. Tibiae completely or predominantly dark; light scaling either as narrow lines or as patches. Tarsal white rings always basal, usually restricted to proximal segments. Claws simple on all legs but with minute spicules at base of external surface. Wing: 10 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Veins usually entirely dark scaled, white scales rarely present on remigium; plume scales restricted to veins Rg, R243, Ry, R3 and M. Abdomen: Tergite I with numer- ous scales; laterotergite with large white or silvery scale patch. Tergites II-VII with broad basolateral silvery patches, progressively more removed from base and extend- ing more tergally on distal segments, on VIII usually merging with its mate middor- sally; basomedian tergal light scaling present or absent on II-VII. Light scaling of sternites varied. FEMALE GENITALIA (fig.16). Deeply retracted into segment VIII, even cerci usually invisible. Segment VIII partially retracted into segment VII, apex visible, nu- merous scales and bristles present; tergite about 0.6 length of tergite VII; sternite about 1.4 of tergite length and about 0.8 of sternite VII. Tergite [X strongly devel- oped, about 0.5 of tergite VIII, undivided, without bristles. Sigma joined to cowl, narrow; continuous with poorly sclerotized insula which bears 2 pairs of setae. Cowl narrow, joined to base of postgenital plate; atrial plates absent. Postgenital plate shorter than cercus, bilobed. Tergite X not developed. Cercus broad, subequal in length to tergite [X or a little longer. Spermathecae 3, 1 a little larger than others. MALES (fig.16). Coloration usually similar to females, sometimes light scales of torus absent (allotecnon) and rarely proboscis with a ventral light patch (whitmorei); sexual dimorphism of head appendages marked. Proboscis usually longer than in fe- males. Palpus porrect, slender, subequal to proboscis; 5-segmented, segments 4 and 5 subequal, narrowed; entirely or predominantly dark scaled, light scales restricted to lower surface of segments 4 and 5; a few short inconspicuous setae on body of seg- ment 4 and apex of segments 3 and 5. Whorls of flagellar segments 1-12 very strong- ly developed, usually with about 40 or more long bristles; flagellar segment 12 and 13 elongate, 12 about 1.25 of 13; torus greatly swollen. Claws of foreleg and midleg enlarged, unequal; larger claw with a blunt submedian tooth; both claws with or without an acute basal external tooth and with external basal spicules more numer- ous than in females, extending on shaft. Hindclaws as in the females. MALE GENITALIA. Segment VIII long and wide, unspecialized. Segment [IX moderately to strongly developed; tergite lobes indistinct or prominent and bearing a few to many moderate to strong setae, connected middorsally by a narrow sclero- tized bridge, sternite strongly developed, usually with distinct median sclerotization bearing several pairs of setae distally. Sidepiece well developed; more or less conical or subcylindrical; mesal membrane developed from base to apex; tergomesal and/or sternomesal margins rarely (Whitmorei Group) with slight median lobes; with numer- ous long bristles and scales laterally and ventrally, tergal surface varied. Claspette rarely completely absent (Whitmorei Group), usually very poorly developed as a small sternal basomesal lobe bearing 1 or rarely 2 thickened apical setae and some- times a few other less differentiated setae. Clasper simple, slender; 1 apical spiniform always present. Aedeagus without teeth; varied in shape, usually large; lateral sclero- tization sometimes produced as a distal horn or lobe. Proctiger strongly developed, prominent; paraproct with a simple, large, heavily sclerotized, curved apical tooth; cercal setae distinct, short. PUPAE. Cephalothorax: Middorsal ridge moderate; all hairs present, variously de- veloped; hair 6-C always smaller than and cephalad of 7-C; hairs 8,9-C always widely separated and far caudad of trumpet base. Trumpet: Not placed on tubercle; varied in length; index from about 3.0 to 9.0; always without tracheoid; pinna short to long. Metanotum: Hairs 10,11-C removed from 12-C, moderately close together. Abdo- men: Hairs 8,10,11-II apparently usually absent; hair 5-II,II] usually caudad of hair 4: hair 1 usually most mesal hair on all segments; hair 9-VIII ventral in position, near Berlin: Subgenus Howardina 11 caudolateral angle or removed considerable distance cephalad; hairs 1-[X,X apparent- ly not developed. Tergite IX strongly developed, its posterior margin smoothly rounded; cercal lobe of female projecting only a short distance beyond genital lobe. Paddle: Variously developed; midrib usually strongly differentiated; marginal spic- ules varied. LARVAE. Head: Width of head capsule subequal to length or slightly greater; la- brum indistinctly differentiated, very narrow; mouthbrushes very numerous, filamen- tous; collar well developed, narrow; ventral part of head capsule (labial plate) rather short; posterior tentorial pit near caudal border; maxillary suture complete, some- times extending caudolaterad of pit; cephalic border of labial plate truncate; aulaeum with filamentous spicules; mental plate well developed. Hair O-C distinct, far re- moved laterad of 1-C; 2-C apparently always absent; 3-C moderately developed; 11, 12,14-C sometimes stellate. Antenna: Short to moderate in length; shaft with or without short spicules. Hair 1-A single to 3-branched. Thorax: Integument glabrous or with more or less distinct spicules. Hairs 0,1,3,4,8,13,14-P, 1,13,14-M, 1,3,4,5,8, 13-T sometimes stellate. Hairs 1-3-P on common tubercle, placed close to middorsal line; tubercles of 5-7-P usually distinctly joined (except in Fulvithorax Section); 9- 12-P far ventrad, poorly developed, none of the hairs distinctly stellate. Hair 5-T always cephalad of hair 3. Abdomen: Integument as on thorax. Hair 12-I never developed, 9-I always ventrad of 7-I; hairs 1,2,5,7,9,13 frequently stellate on seg- ments I-VI; hair 6-I-VI always well developed, long; hair 7-I long. Segment VIIT: Comb scales in a single regular row to an irregular patch of 3-5 rows. Siphon: Index varied from 2.0-9.0. Integument smooth, imbricate or spiculate. Pecten in a straight line, usually at least reaching to middle of siphon. Hair 1-S usually near middle; ac- cessory hairs not developed. Acus varied or rarely absent (walkeri). Anal Segment: Saddle always incomplete; integument smooth, imbricate or spiculate; caudal margin with or without spines; acus absent. Ventral brush with 5 or 6 pairs of hairs, on a more or less distinctly sclerotized boss, with or without distinct grid bars. DISCUSSION. For a general discussion of taxonomic characters, bionomics, dis- tribution, systematics and affinities see the preceding chapter entitled “General Con- siderations.” The subgenus Soperia is here synonymized with Howardina. The taxon involved represents the Whitmorei Group of 2 species in the Sexlineatus Section, character- ized primarily by the total absence of the claspette in the male genitalia. In oth- er respects the Whitmorei Group obviously belongs in this section of Howardina and it is evident that there is a tendency within this subgenus to reduction in the clasp- ette. Further, the name Soperia cannot be applied to the Whitmorei Group without recourse to the International Commission on Zoological Nomenclature since the nominal type species, dominicii, designated by Komp is a synonym of sexlineatus which obviously belongs to a different taxonomic group. In view of the above-men- tioned taxonomic interpretation, there is no need at this time to request the Com- mission to vary the type species of Soperia, and this subgenus becomes a subjective synonym of Howardina by strict application of the rules in the current code. KEYS TO SECTIONS ADULTS & Prescutellar space with single median golden, yellow or silvery line; parater- gite without scales ob ee ee WAL ARERI SECTION 12 2(1). 2(1). 3(2). 4(3). 21 y:, IL), Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Prescutellar space with paired lateral hie yellow, golden or silvery lines; paratereideowmitivscdies ig a ee ee eee ee ee Mesonotum with acrostichal line always dark . . SEXLINEATUS SECTION Mesonotum with at least acrostichal line yellow or golden, sometimes anteri- or half or more of mesonotum entirely or largely yellow or golden-scaled FULVITHORAX SECTION MALE GENITALIA Tergal surface of sidepiece with a very dense vestiture of large bristles ob- scuring claspette area from dorsal aspect (15. bahamensis) . . . in part WALKERI SECTION Tergal surface of sidepiece with sparse ‘vestiture of short or moderate bristles MOL Goscuring claspeite area trom dorsal ASPeCt an... ek ce, ee ee Claspette completely absent Eee Group) |. e . in part SEXLINEATUS SECT ION Claspette represented by1 more or less strongly developed sternomesal lobe or plaque bearing usually 1 thickened apical bristle, rarely 2 ormore . . .3 Sidepieces separated from each other at base sternally by a distance less than maximum width of aedeagus . ... . . .SEXLINEATUS SECTION Sidepieces separated from each other at base Noe by a distance greater LAall WiAxteI Wicd IY OL. acccaeus 2... ge eae cate ow do cone ee Claspettes connected across midline by more or less distinct sclerotized bar Ae Sere Oe eer ae eer FULVITHORAX SECTION Claspettes not connected across midline . . . . . . WALKERI SECTION PUPAE Hair 5-C longer than distance between its base and that of trumpet, always double; hair 2-[V-VI usually within basal 0.66 of itstergite. . . . SEXLINEATUS SECTION Hair 5-C shorter than distance bétween its base and that of trumpet, varied in branching; hair 2-IV-VI usually distad of basal 0.75 ofits tergite. . . .2 Hair 9-VI minute, less than 0.5 of hair 2-VI. . FULVITHORAX SECTION Hair 9-VI subequal to or longer than hair 2-VI. . . . WALKERI SECTION LARVAE Caudal border of saddle without distinct marginal spicules; comb scales al- ways in a patch of 3 or 4irregularrows . . FULVITHORAX SECTION Caudal border of saddle with conspicuous long marginal spicules; comb scales usually in a single even row, rarely in a patch of 2-4 irregular rows (J6. | Hair 13-P notdeveloped ........... .. . .WALKERI SECTION Oe eee er ee OE ete ace hd cia a oe do sane beech: Ga Berlin: Subgenus Howardina 13 3(2). Hair 9-P poorly developed, single (Ioliota Group). . coo Ieper WALKERI SECTION Hair 9-P strongly developed, at Jeast A-branched . SEXLINEATUS SECTION FULVITHORAX SECTION FEMALES. Medium sized species, wing 2.8-3.6 mm. Head: Scales predominantly or entirely light, submedian and lateral dark patches sometimes large. Erect scales present on disc of vertex. Torus yellowish to dark brown, with a few moderately broad dark scales mesally. Palpus entirely dark scaled. Thorax: Mesonotum varying from nearly entirely golden or yellow to predominantly dark bronzy but with at. least golden acrostichal line developed; median prescutellar light line not developed. Paratergite with scales. Acrostichal bristles absent on disc. Pleural scales silvery or dingy golden; ppn, psp, and ssp with scales; most of mep covered with large continu- ous scale patch. Legs: Midfemur and hindfemur without distinct distal light spot on anterior surface. Apex of tibia and base of tarsal segment 1 of all legs with white scales; all other tarsal segments dark. Wing: Veins entirely dark scaled. Abdomen: Sternites II-[V without dark scales. MALES. Larger claw of foreleg and midleg with submedian tooth only. MALE GENITALIA. Lobes of tergite [IX indistinct to moderately prominent, with numerous strong setae. Sidepieces separated from each other at base by a distance greater than diameter of aedeagus at that level; tergomesal and sternomesal margins without any indication of lobes. Claspette prominent, connected at base with its mate by more or less distinct sclerotized bar; apical and subapical bristles varied in development. PUPAE. Cephalothorax: Hair 5-C distinctly shorter than distance between its base and that of trumpet. Abdomen: Hair 5-IV,V usually single, extending beyond apex of tergite following; 2-IV-VI usually distad of 0.75 of its tergite; 9-II-VI cephalad of hair 6; 9-VII moderate to strong, close to hair 6, not removed cephalad. LARVAE. Stellate hairs moderately developed. Antenna: Short; without spicules; hair 1-A single. Thorax: Integument glabrous; hairs 5-7-P on separate tubercles; hair 9-P usually single or double; hair 13-P absent. Abdomen: Integument glabrous; hairs 2,5,9-II-VII usually stellate, with few branches; hair 8-VII cephalodorsad of 9-VII. Segment VIII: Hair 5 stellate. Comb scales always in a patch of 3,4 irregular rows; individual scales rounded apically. Siphon: Index 2.2-2.3. Integument glabrous; pig- mentation very strong. Acus distinct. Pecten teeth with basal denticles. Anal Segment: Saddle without marginal spicules. Hair 1-X short to moderate, double or triple; hair 3-X single, 2-X multiple Ventral brush with 6 pairs of hairs arising from a strongly sclerotized boss. DISCUSSION. The Fulvithorax Section is probably the most generalized lineage in Howardina and is strongly differentiated in all stages from all the other sections of the subgenus. The pattern of ‘ornamentation of the mesonotum of the adults is unique in the subgenus and shows considerable variation whereas it is stabilized in the other sections. The male genitalia are readily recognized by the widely spaced side- pieces and the relatively large prominent claspette. The larvae are immediately sepa- rated from all the other sections by the characteristic comb patch and the absence of caudal marginal spicules on the anal saddle. The pupal stage has the simplest chaeto- taxy in the limited branching and the relatively small size of the more conspicuous hairs. In nearly all these characters, the Fulvithorax Section shows the closest simi- 14 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 larity to the subgenus Finlaya from which Howardina was probably derived. Further- more, as in typical American Finlaya, the natural breeding sites of the Fulvithorax Section are treeholes and broken bamboo. The Fulvithorax Section presents a challenging taxonomic problem, which, unfor- tunately, I have not been able to resolve satisfactorily because of the absence of reared associated material from nearly all the critical areas of its wide reported distri- bution (fig.2) which is largely confined to the Atlantic drainages and extends from Nicaragua to southern Brazil. Formerly, 3 species with nearly overlapping distribu- tions were recognized almost entirely on the basis of the extent and arrangement of the golden scaling of the mesonotum. The name fulvithorax was applied to forms with the mesonotum nearly entirely golden, septemstriatus to specimens with dis- crete lines of golden scales separated by dark scales, and arborealis to specimens in- termediate in this respect. This situation suggested that only 1 very variable species was involved. However, closer analysis of the adults available for the present study and the examination of associated reared immature stages from 3 areas reveals a much more complex situation. In the present study, I am recognizing only 1 group with 5 different one in the Fulvithorax Section: fulvithorax, with a wide distribution from Venezuela and Trini- dad to southern Brazil and occurring a considerable distance inland in Brazil and Peru; septemstriatus, restricted to lowland and moderate elevations in Nicaragua, Panama and Colombia; arborealis, from the Guianas and the lower Amazon; aureo- lineatus, primarily from the state of Bahia but apparently extending north to Para and south to Rio de Janeiro; and the unnamed Iquitos form from Peru which may be an inland type of arborealis. A. fulvithorax, septemstriatus and arborealis all show more or less distinct north-south clines in the extent of mesonotal golden scales, the southern populations being in general darker. In the case of fulvithorax, the inland populations also tend to be darker than the coastal. In any given area, only 2 of these forms appear to be present, except that 3 may occur in Para, and in all instances the different sympatric forms are quite distinct, without any intermediates. This suggests that distinct species are involved. However, the material available for this study is very limited and it is possible that the 5 forms recognized here are only subspecifical- ly distinct and that there might be intergradations between them. It is even possible that only 1 very variable species is involved. The problem cannot be definitely re- solved without progeny and individual rearings from areas of contact and overlap. KEYS TO SPECIES ADULTS ]. Mesonotum usually entirely golden-scaled from anterior promontory to wing root, rarely a few darker scales along dorsocentral lines posteriorly; no corispicuous dark bronzy scalesonfossa. ... . . .. . JL. fulvithorax Mesonotum not entirely golden-scaled in anterior portion, with at least some conspicuous dark bronzy scales on fossa, usually with a more or less con- spicuous linear pattern of golden scales surrounded by dark bronzy scales . ae 2(1). Central part of anterior mesonotum with at least anterior inner dorsocentral lines golden in addition to the acrostichal line. . .. co - Central part of anterior mesonotum with only the acrostichal line golden .4 Berlin: Subgenus Howardina 15 3(2). Lateral mesonotal light line conspicuous anteriorly, broad, not interrupted at or before scutal angle . . . . . . 2. Septemstriatus Lateral mesonotal light line either completely absent Or very narrow or inter- rupted at scutal angie; es ee ee eras 4(2). Posterior outer dorsocentral golden line distinct, linear . . 4. aureolineatus Posterior outer dorsocentral golden line represented by a patch caudad of scuta anes 68 bee ee ae ae ae oe ee ee MALE GENITALIA (5. Iquitos form unknown) © F; Subapical seta of claspette usually less than 0.7 of length of apical, latter not reaching middle of sternomesal margin of sidepiece . Die: . . 1. fulvithorax; 2. septemstriatus Subapical seta of claspette usually subequal i in length to apical, latter extend- ing beyond level of middle of sternomesal margin of sidepiece. . . . .2 2(1). Several subapical setae of claspette distinctly thickened . . 4. aureolineatus None of the subapical setae of claspette thickened . . . . . 3. arborealis PUPAE (4. aureolineatus and 5. Iquitos form unknown) iP Hair 1-II usually with 4 or more branches. . 1. fulvithorax; 2. septemstriatus Haiw-i ll usually double @ 66456 sha a6 2 we ees ie Ay reals LARVAE (4. aureolineatus and 5. Iquitos form unknown) L Hair 6-C usually double; 11-C double; most pecten teeth without basal den- telee 2 ess . . 3. arborealis Hair 6-C single; 11-C with more than 2 branches; all pecten teeth with basal Genes: (ilies i ere ee ae ee TO a eee: 2(1). Hair O-P long, usually 14-branched (10-21); gills moderate, ha, unequal . . 1. fulvithorax Hair 0-P short, usually $-branched (4 0 lls long slendet and subequal. SORE Oe BO8 .2. septemstriatus 1. Aedes (Howardina) fulvithorax (Lutz) Figs.2,8,10,11 1904.Haemagogus fulvithorax Lutz, 1904:47,66,73. TYPE: Holotype ?, bridge Ipe-Arcado (Goias), Brazil, date and collector not specified [LU]. 1906.Taeniorhynchus palliatus Coquillett, 1901a:61. TYPE: Holotype , Trinidad, locality and date not specified, F.W. Urich [USNM, 9140]. Synonymy with fulvithorax by Howard, Dyar and Knab (1917:844). 1947.Aedes (Howardina) tachirensis Anduze, 1947:355-357. TYPE: Holotype ?, Las Mesas de 16 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Seboruco, Dto. Jauregui (Tachira), Venezuela, date and collector not specified [A] . Synony- my with fulvithorax by Lane (1951:335). Aedes (Howardina) fulvithorax of Bonne and Bonne-Wepster (1925:373-375); Dyar (1928:228- 229); Shannon (1931a:8,23,24,25; 1931b:142,143,146,161); Edwards (1932:155); Soper, Pen- na et al (1933:576); Lane (1936b:186); Whitman and Antunes (1937:819-821); Lane (1939: 99); Floch and Abonnenc (1942a:5; 1942b:6); Anduze (1947:355); Lane (1951:335; 1953: 706-708); Horsfall (1955:413,471,483); Martinez and Prosen (1955:29-31); Vargas (1956:22); Stone, Knight and Starcke (1959:174); Fauran (1961:29); Prosen, Carcavallo and Martinez (1964:104); Forattini (1965:400); Cova Garcia, Sutil and Rausseo (1966a:57,285; 1966b:11, 73,141 ,169,333-334). Aedes fulvithorax of Howard, Dyar and Knab (1912:65); Urich (1913:528); Howard, Dyar and Knab (1917:844-846); Dyar (1918:73,80); Costa Lima (1930:255); Davis and Shannon (1931: 716,717,718); Martini (1931:206-208); Lane (1936a:131); Causey and Kumm (1948:475,476, 477); Causey and Santos (1949:478); Causey, Kumm and Laemmert (1950:302,307); Matheson (1950:325); Delponte, Castro and Garcia (1951:239,240); Whitman (1951:263); Floch and Fauran (1954:48); Aitken (1960:4); Belem Virus Lab (1964:101,103); Bates (1949:205,320). Gualteria fulvithorax of Theobald (1907:552); Peryassu (1908:45,179-180); Theobald (1910: oe Haemagogus fulvithorax of Lutz (1905:67); Dyar and Knab (1906a: 166,167). 3 Taeniorhynchus palliatus of Coquillett (1906b:24); Theobald (1910:432). FEMALE. Wing: 2.97 mm. Proboscis: 2.34 mm. Forefemur: 2.16mm. Abdomen: about 2.88 mm. Head: Scaling predominantly golden, a few dark scales present but not forming distinct lateral patches; narrow median scales occupying an extensive tri- angular area extending in the back to the outer erect scales and wide in front. Palpus about 0.19 of proboscis. Torus light brown to yellow, usually darkened on mesal sur- face. Thorax: Mesonotal integument yellowish to light brown; scaling usually entire- ly golden to just before level of prescutellar space; numerous linear dark bronzy scales along prescutellar rows, sometimes extending forward along dorsocentral bris- tles, a few similar dark scales in front of prescutellar space, a variable patch of similar dark scales above supraalar bristles, no dark scales on fossa; golden scales at scutal angle not markedly broadened (fig.8). Median scutellar lobe with numerous broad dark scales and a few golden in middle from base to apex; lateral lobes predominant- ly with narrow golden scales. Pleural scaling silvery, with slight yellowish tinge; apn predominantly with narrow scales, a few moderately broadened ones on lower and posterior parts; ppn with sparse vestiture of short linear golden scales, usually with only a few broadened scales in lower part. Abdomen: Median basal light patches usu- ally present on tergites III, IV, light brown or golden; always present on V,VI, usually with slight yellowish tinge. MALE. Wing: 2.36 mm. Proboscis: 2.1 mm. Forefemur: 1.49 mm. Essentially sim- ilar to female but lighter in color, dark scales of prescutellar area and midlobe of scu- tellum less numerous; apn with large patch of broad scales on lower posterior sur- face. Palpus distinctly shorter than proboscis. MALE GENITALIA (fig.10). As figured; diagnostic characters as in the key. Ninth tergite lobe not prominent, with about 5 bristles. Claspette with subapical seta not thickened, less than 0.7 of apical, latter not reaching middle of sternomesal margin of sidepiece. Clasper with 1,2 subapical hairs. PUPA (fig.10). Abdomen: 3.24 mm. Trumpet: 0.38 mm; index 4.5. Paddle: 0.73 mm. Chaetotaxy as figured; hairs moderately to darkly pigmented. Apparently indis- tinguishable from septemstriatus except perhaps by the longer trumpet. Readily sep- arated from arborealis by hair 9-VI being minute, less than 0.5 of hair 2-VI, and more numerous branches in hairs 1-II, 9-VII, VIII. Cephalothorax: Mesonotum moderately Berlin: Subgenus Howardina 17 pigmented between trumpets; lateral areas and metanotum slightly darker. Hair 1(1, 1+2),° 201); 301); 4Q)s- 5Qa1-3),- 601), 7; 1-3)380- F3), 9Uy 10, 2-4) 11), 12(2, 1-4). Abdomen: Tergite I deeply pigmented except for usual membranous ar- eas; tergite II uniformly moderately pigmented; tergites and sternites HI,[V with moderate pigmentation but anterior border darker; rest of segments lighter. Segment I: hair 1(10-14 primary branches, each with many secondary ones), 2(2, 1-4), 3(1), 4(4, 2-6), 5(3, 2-5), 6(1), 7(2, 1-3), 9(1), 10(1, present on majority of specimens but sometimes absent on one side). Segment II: hair 0(1), 1(7, 4-13), 21, 1-2), 31, 1-3), 4(4, 3-5), 5(1, 1-2), 6(1, 1-2), 7(1, 1-4), 9(1). Segment III: hair 0(1), 1(2, 1-4), 2(1), 3(1)s.435 1-4; 501 2: 1-2),.601), 78:2-4).8@, 13), 90), 100, 1-2); LU, 140). Segment IV: hair 0(1), 1(1, 1-2), 2(1), 3(3, 1-4), 4(2, 1-3), 5(1), 6(1), 7(3, 2-3), 8@, 2-3), 9(1), 10(1, 1-3), 11(1), 14(1). Segment V: hair 0(1), 11, 1-2), 2(1), 31, 1-2), 4(2, 1-4), 5(1), 6(1), 7(3, 2-4), 8(2, 1-3), 9(1), 10C1, 1-2), 111), 141). Segment VI: hair O(1, 1-2), 1(1), 2(1), 3(1), 4, 1-2), 51), 6(1), 71, 1-2), 8@2, 1-2), 9(1), 101), 11(1), 14(1). Segment VII: hair 0(1), 1(1), 21), 3), 41), 5(1), 6(@, 1-4), 70), 8@, 1-3), 9(4, 3-5), 10(1), 111), 14(1). Segment VIII: hair 0(1), 4(1), 9(7, 5-8), 14(1). Paddle: Uniformly lightly pigmented, midrib darker; external margin slightly ser- rated; hair 1-P slender; male genital lobe extending to about 0.37 of paddle, female genital lobe to 0.29. LARVA (fig.11). Head: 0.89 mm. Siphon: 0.7 mm. Anal Saddle: 0.27 mm. Both hairy and nonhairy forms known. Very similar to septemstriatus from which it can be separated apparently by the longer, more profusely branched hair 0-P and the broad, short unequal gills. Head: Pigmentation moderate except around mouthparts where it is deeper; integument smooth. Mental plate with a strong median tooth and 11(11,12) distinct teeth on either side, often a smaller detached tooth at base. Hair O(1), 1(1), 3(1), 4(13, 10-16), 5(1), 6(1; moderate), 7(3, 2-4), 8(1), 9(2, 1-2), 10(1), 11(12, 11-16; stellate), 12(3, 2-4), 13(1), 14(4, 3-8), 15@, 2-4). Antenna: Length about 0.48 of head; pigmentation deeper than on head capsule; all hairs single; hair | at 0.5 from base. Thorax: Apices of long hairs attenuate; barbs present on long and stellate hairs. Prothorax: hair 0(14, 10-21), 1(4, 3-4), 2(1), 3(7, 4-8), 444, 4-9), 5@, 2-4), 6(1), 7(3, 2-3), 8(11, 8-18), 9(2, 1-5), 10(1), 11@, 1-4), 121), 14(21, 17-28). Mesothorax: hair 1(5, 3-8), 2(1), 3(1), 4(1), 5(1), 6(5, 4-6), 7(1), 865, 4-7), 965, 4-7), 10(1), 11(2, 1-3), 12(1), 13(15, 12-23), 14(9, 8-16). Metathorax: hair 1(5, 4-12), 2(1), 3(4, 3-6), 4(6, 5-13), 5(5, 4-13), 6(1), 7(5, 4-8), 809, 8-22), 9(4, 3-5), 10(1), 11(1, 1-2), 12(1), 13(11, 8-18). Abdomen: Stellate hairs with branches of unequal length. Segment I: hair 1(8, 7-13), 2(10, 4-10), 3(1), 4(6, 5-6), 5(12, 10-13), 6(4, 3-4), 7€1, 1-2), 9(6, 5-8), 10(1), 11(12, 6-13), 13(8, 5-12). Segment II: hair 0(1), 1(1:0;:3-441); 209; 6-10), 31), 465, 3-8), 5(10,7-11); 66, 3-4), 73, 2-7), (1), 9(8, 5-10), 10(1), 111), 12(1), 13(8, 5-14). Segment III: hair 0(1), 1(4, 2-8), 2(7, 7-9), 3(1), 4(1), 5(9, 6-12), 6(2), 7(6, 5-10), 8(1), 9(9, 6-13), 10(1), 11(1), 12(1), 13, 4-12), 14(1). Segment IV: hair 0(1), 1(5, 2-9), 2(8, 7-9), 3(1), 4(1), 5(9, 6-11), 6(2), 7(5, 5-8), 8(1), 9(8, 7-11), 10(1), 11(1), 121), 134, 2-10), 14(1). Segment V: hair 0(1), 1(3, 2-9), 2(7, 6-9), 3(2, 1-2), 4(1), 5(6, 6-9), 6(2), 7(5, 5-8), 8(1), 9(8, 6-10), 10(1), 1101), 12(1), 13(5, 2-11), 14(1). Segment VI: hair 0(1), 1(5, 2-10), 2(7, 6-9), 3(1), 4(1), 5(7, 4-11), 6(1, 1-2), 7(4, 3-7), 8(1, 1-2), 9(4, 4-11), 10(1), 11€1), 121), 13(6, 5-13), 14(1). Segment VII: hair 0(1), 1(3, 2-8), 2(7, 6-10), 3(1), 4(1), 5(4, 3-8), 6(3, 2-3), 7(1), 8(5, 4-7), 9(4, 3-6), 10(1), 11(1), 121), 134, 3-10), 14(1). Segment VII. Comb scales 20(19-24), in a patch of 3 irregular rows; individual scale elongate, rounded apically and evenly fringed with short spinules; hair 0(1), 1(5, 4-10), 2(1), 3(7, 5-8), 4(1), 5(6, 5-11), 14(1). Siphon: Index about 2.25-2.3; darkly pigmented 18 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 and with imbricate sculpturing throughout. Pecten extending to about 0.34-0.38; with 13 teeth (10-15), usually all with 1-3 basal denticles, terminal tooth not mark- edly enlarged. Hair 1(3, 3-4), 2(1), 3-5 not visible, 6(1), 7(1), 8(@2, 1-3), 9(1). Anal Segment: Saddle darkly pigmented, with uniform imbricate sculpturing. Gills broad and short, unequal, ventral about 0.5 of dorsal. Hair 1(3, 2-7), 2(4, 3-5), 3(1), 4a(4, 3-5), 4b(3, 3-4), 4c(3), 4d(3, 3-4), 4e(3, 3-4), 4f(3, 2-4). SYSTEMATICS. Adults of fulvithorax show considerable variation in thoracic or- namentation in different areas of the reported wide distribution of this species but are readily separated from the other species of the group by the much greater amount of golden scaling on the mesonotum in both males and females. The above descrip- tions of all stages are based entirely on material from Trinidad, the type locality of palliatus. The single specimen I have seen from Venezuela, a female paratype of tach- irensis, is indistinguishable from the Trinidad material. In these 2 northern popula- tions, the integument of the thorax is light yellowish tan and the mesonotal scaling is bright yellow golden without any indication of dark scales on the fossa or anterior dorsocentral lines. The Surinam specimen agrees with this description also. The French Guiana material is similar in scaling but the thoracic integument is brown. As one proceeds south along the Atlantic lowlands of Brazil (Para, Bahia and Rio de Ja- neiro) the mesonotal golden scaling becomes darker, more coppery and the integu- ment tends to become darker brown; however, in all these populations the fossa is without dark bronzy scales. At higher elevations in Brazil (Sao Paulo, Goias, Matto Grosso), the darkening of the integument and scaling becomes even stronger and there is a tendency for the appearance of a few dark bronzy scales in the fossa and along the posterior dorsocentral line. The single female from Iquitos resembles the interior populations of Brazil but has more extensive dark-scaled patches on the head than any other one I have seen. In the scanty material of immature stages available from populations other than palliatus, no striking differences are apparent. No clear-cut separation can be made of fulvithorax from septemstriatus at this time in the pupal stage but the larvae of the 2 species are apparently distinct on the _ characters given in the key and diagnoses. BIONOMICS. The immature stages of fulvithorax have been found only in tree- holes. The adults are primarily diurnal in activity but a few specimens have been col- lected at night. Females feed readily on experimental animals in the laboratory. They have been investigated as possible vectors of yellow fever virus (Davis and Shannon 1931; Ait- ken 1960). The virus will survive and apparently develop in these mosquitoes as shown by successful infection of rhesus monkeys by inoculation of experimentally infected fulvithorax but all attempts to transmit the virus by bite have been negative. DISTRIBUTION (fig.2). Lowlands to moderate elevations in Venezuela (Maracai- bo and Orinoco drainages), Trinidad, Guyana, Surinam, French Guiana, Brazil (Ama- zon, Paraguay and Parana drainages, and states of Espirito Santo and Rio de Janeiro) and Peru (Amazon drainage). Material examined: 152 specimens; 20 6, 62 9, 29 pu- pae, 41 larvae; 30 individual rearings (4 pupal, 20 larval, 6 incomplete). BRAZIL. Bahia: Piraja, 2 Apr 1929, [C. Davis and] R.C. Shannon, 2 Ip, 2 6, 1 6 genitalia, 5 L, 6 1; 1938, 1 6d; 1 1 [USNM]. Espirito Santo: Santa Teresa, Apr 1940, 2 ° (Martinez and Prosen, 1955:31; MEPRA). Valle do Chanaan, 173 @ (Davis and Shannon 1931:715; Whitman and An- tunes 1937:819-820). Goias: Anapolis, Sept 1936, 1 6, 1 9 [USNM]. Matto Grosso: Guiaba, Feb 1935, 2 2; Mar 1935, 3 ? [USNM]. Maracaju, July 1937, 1 9; Nov 1937, 1 9; Dec 1937, 8 9; 3 2 [USNM]. Pocinho (Lane 1936b:186). Minas Gerais: Passos, 1945-47, 23A (Causey and Kumm Berlin: Subgenus Howardina 19 1948:475,477); 1840A (Causey, Kumm and Laemmert 1950:302,307); 386A (Causey and Santos 1949:478). Para: Belem, IAN forest, 105 9; IAN and Utinga forests, 9 2; Utinga forest, 9 ? (Belem Lab Report 1964:101,103,109,114). Belterra, June 1938, 1 2 [USNM]. Ilha de Marajo, 126 ? (Kumm and Novis 1938:502). Urucurituba, 18 Mar, C.H. Townsend, 1 9 [USNM] . Rio de Janeiro: Apr 1938, 4 9 [USNM]. Sao Paulo: Guariroba; Nhandeara; Nipoan; Planalto; Villa Poloni (Lane 1936a:131). State not specified: 1 3,1 [USNM]. FRENCH GUIANA. Baduel (Fauran 1961:29). Balourou, Crique Anguille, Galion (Floch and Abonnenc 1947:10). Placer Isnard, June 1940, E. Abonnenc, 1 @ [UCLA] (Floch and Abonnenc 1947:10; Fauran 1961:29). Rorota, 8 May 1941, E. Abonnenc, 1 ¢ [UCLA] (Floch and Abon- nenc 1947:10). Souvenir (Floch and Abonnenc 1947:10). Locality not specified, 1944, 1 d, 1 [USNM]. | GUYANA. Mazaruni, 25 June 1936, 1 2 [USNM]; 6 July 1936, Leslie, 1 d (KO 208A-1) [UCLA]. PERU. Iquitos, Mar-Apr 1931, R.C. Shannon, 1 9 [USNM]. SURINAM. Moengo, 19 Mar 1946, H.H. Stage, 1 L [USNM]. Locality not specified, 1 (36- 69) [ITH]. TRINIDAD. Nariva: Charuma forest, 27 Aug 1964, treehole, A. Guerra, 1 lp? (TR 634-108) [UCLA]. St. Andrew: Mt. Harris, elev. 250 ft, 16 July 1964, treehole, F. Powdhar (TR 566), 1 Ipd (566-102), 1 Ip? (566-108) [UCLA]. St. George: Arima, Aripo Valley, elev. 500 ft, 16 Apr 1964, bamboo, A. Guerra (TR 319), 1 lpd (319-148), 2 Ip? (319-139,149), 1 p? (319-111), 2 IP (319- 128,137), 1 L; Cumaca road, elev. 500 ft, 14 Jan 1965, A. Guerra, 1 2 (TR 943); Verdant Vale, elev. 1000 ft, 10 Sept 1964, treehole, A. Guerra, 2 lp? (TR 680-102,103) [UCLA]. Monos Island, Grand Fond, 17 May 1964, treehole, A. Guerra (TR 410), 2 Ipd (410-106,107), 1 Ip? (410-111); 1 Aug 1964, bamboo, R. Martinez (TR 584), 1 lp? (584-11), 1 pd (584-102); 2 Aug 1964, bam- boo, R. Martinez (TR 591), 1 lpd (591-109), 4 lp? (591-113,114,119,121), 1 pd (591-108), 1 pe (591-107), 1 L [UCLA]; 17 Nov 1905, F.W. Urich, 3 6, 1 ?; June 1906, 2 Ip? (20-2,3), 1 12 (20-1), 1 lp (20-4), 4 9? [USNM]. San Rafael, Arena forest, Oct 1965, 2 9 (TR 1432), Dec 1965,3 9 (TR 1441) [UCLA]. Port of Spain, 9 Sept, 1 9; 1 9 [USNM] . County not specified: 26 Aug 1945, 1 Ip; 9 Sept 1945, 1 ? [USNM]. VENEZUELA. Bolivar: Distrito Heres (Anduze 1947:355). Rio Orocopiche (Cova Garcia, Su- til and Rausseo 1966b:73). Tachira: Chururu, Distrito Uribante (Anduze 1947:357). Las Mesas de Seboruco, Distrito Jauregui, June 1944, P.J. Anduze, 1 9 [USNM] (Anduze 1947:357; Cova Garcia, Sutil and Rausseo 1966b:73). 2. Aedes (Howardina) septemstriatus Dyar & Knab Figs.2,8,12,13 1907.Aedes septemstriatus Dyar and Knab, 1907:10. TYPE: Holotype ? (8), Bluefields, Nicaragua, W.F. Thornton [USNM, 10144]. Aedes (Howardina) septemstriatus of Bonne and Bonne-Wepster (1925 :366-367); Dyar (1925:141, 148; 1928:232-233); Shannon (1931:142,147); Edwards (1932:155); Lane (1939: 100); Floch and Abonnenc (1942a:6; 1942b:5; 1944:4; 1947:10); Martinez and Prosen (1955:27-29); Reyes (1955:74,89); Trapido, Galindo and Carpenter (1955:530,537,538); Trapido and Galindo (1957:122,125); Stone, Knight and Starcke (1959:174); Fauran (1961:29); Belkin, Schick and Heinemann (1965:42); Prosen (1965:42); Forattini (1965:239,242,403-404). Aedes septemstriatus of Theobald (1910:485); Howard, Dyar and Knab (1917:846-847); Dyar (1918:73); Komp (1936:63); Antunes (1937a:16; 1937b:76); Arnett (1949:238); Galindo, Tra- pido and Carpenter (1950:102,104,105 108,112,113); Carpenter and Peyton (1952:681); Galin- do, Carpenter and Trapido (1955:163); Reyes (1955:75); Komp (1956:39); Stone and Knight (1956:225); Blanton and Peyton (1958:31-32). Aedes arborealis of Antunes (1937b:76); Reyes (1955:74,89); Boshell-Manrique and Osorno-Mesa (1944:177). 20 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 FEMALE. Wing: 3.15 mm. Proboscis: 2.53 mm. Forefemur: 2.01 mm. Abdomen: about 2.8 mm. Head: A large patch of broad dark scales on each side of a narrow me- dian stripe of narrow golden scales, this stripe narrowed caudad. Palpus about 0.14 of proboscis. Torus dark brown. Thorax (fig.8): Mesonotal integument dark brown; mesonotum with a striking pattern of golden lines on a field of dark bronzy scales, latter sometimes with a few scattered golden scales; golden scaled lines as follows, narrow acrostichal bifurcating at prescutellar space, narrow inner dorsocentral not always reaching prescutellar space, outer dorsocentral arising at about level of scutal angle where it is broad and extending as a narrow line to scutellum; lateral marginal line broad, golden, interrupted in front of wing root, golden scales at scutal angle broader than in fulvithorax; fossa predominantly with dark bronzy scales, but some- times with some scattered golden scales; space dorsad of supraalars with a large con- spicuous patch of dark bronzy scales. Scutellar ornamentation as in fulvithorax. Pleu- ral scaling denser and more silvery than in fulvithorax; scales of apn predominantly broad and very dense; ppn scale patch more extensive than in fulvithorax and with distinct broad scales in lower part. Abdomen: Median basal tergal light patches essen- tially as in fulvithorax except that scales on V,VI more silvery, without distinct yel- low tinge. MALE. Wing: 2.7 mm. Proboscis: 2.57 mm. Forefemur: 1.84 mm. Mesonotal or- namentation markedly different from female; anterior part predominantly with gold- en scales as in fulvithorax except for very narrow inconspicuous lines of dark bronzy scales on anterior dorsocentral lines and on each side of moderately broad acrostichal line, fossa usually without any bronzy scales; posterior part of mesonotum with more extensive pale scaling than in female but upper supraalar area with a distinct large patch of bronzy scales. Head with smaller dark patches than in female; palpus longer than in fulvithorax, subequal in length to proboscis. MALE GENITALIA (fig.12). As figured; diagnostic characters as in the key. Ex- tremely similar to fulvithorax but apparently with a more prominent IX tergite lobe with 8,9 bristles, and clasper with 2,3 longer subapical hairs. PUPA (fig.12). Abdomen: 2.97 mm. Trumpet: 0.48 mm; index 4.3-4.5. Paddle: 0.76 mm. Only nonhairy form known. Extremely similar to and apparently indistin- guishable from fulvithorax ; separated from arborealis by the same characters as fulvi- thorax. Chaetotaxy as given below; hairs moderately to darkly pigmented. Cephalo- thorax: Mesonotum lightly pigmented; metanotum moderately pigmented; trumpet deeply pigmented and sharply contrasting. Hair 1(1), 2(1, 1-2), 3(1), 4C1, 1-2), 5(2, 1-3), 6(1), 7@2, 1-2), 8(1, 1-2), 9(1), 10(2, 1-3), 11(1), 12(2, 1-3). Abdomen: All ab- dominal segments lightly pigmented, first 3 segments slightly darker; anteromedian area of sternites III and IV with darker patches; integument imbricate. Segment I: hair 1(9-15 primary branches subequal in length and each with many secondary branches), 2(1), 3(1), 4(5, 2-6), 5(3, 2-5), 6(1), 7(1), 9(1), 10(1), and 11(1, found occasionally). Segment II: hair 0(1), 1(5, 2-7), 2(1), 3(1), 4(4, 2-6), 5(1, 1-2), 6(1), 7(1, 1-2), 8(1, found occasionally), 9(1). Segment III: hair 0(1), 1(1, 1-2), 2(1), 3(1), 4(2, 1-3), 5(1), 6(1), 7(3, 1-4), 8(2, 1-3), 9(1), 10(1, 1-2), 11(1), 14(1). Segment IV: hair O(1), 1(1), 2(1), 3@, 1-3), 4(1, 1-2), 5(1), 6(1), 7(@2, 1-3), 8(@2, 1-3), 9(1), 10C1, 1-2), 11(1), 14(1). Segment V: hair 0(1), 1(1), 2(1), 3(1, 1-2), 4(2, 1-3), 5(1), 6(1), 7(3, 1-4), 8(2, 1-2), 9(1), 10(1), 11€1), 14(1). Segment VI: hair 0(1), 1(1), 2(1), 3(1), 4(1), 5(1), 6(1), 701), 8(2, 1-2), 9(1), 101), 11(1), 14(1). Segment VII: hair 0(1), I(1), 21), 3(1), 41), 51), 6(1, 1-2), 7(1), 8(2, 1-2), 9(3, 2-4), 10(1), 11(1), 14(1). Segment VIII: hair 0(1), 4(1), 9(5, 4-6), 14(1). Paddle: Lightly pigmented, midrib slightly darker; apex not emarginated and with minute denticles; hair 1(1, 1-2). Male Berlin: Subgenus Howardina ) 21 genital lobe extending to 0.35 of paddle, female genital lobe to 0.31. LARVA (fig.13). Head: 0.9 mm. Siphon: 0.78 mm. Anal Saddle: 0.32 mm. Only nonhairy form known. Very similar to fulvithorax but apparently distinct from the latter in the short hair O-P with few branches and gills long, slender and subequal. Chaetotaxy as given below; hairs moderately pigmented. Head: Pigmentation moder- ate except for areas around mouthparts where it is deeper. Mental plate with a strong median tooth and 12 distinct teeth (10-12) on each side, often a smaller detached tooth at base. Hair 0(1). 1(1), 3(1), 4(15, 9-16), 5(1), 6(1), 7(4, 2-6), 8(1), 9(1), 10(1), 116, 3-7), 12(2, 1-3), 13(1), 14@2, 2-3), 15(2, 1-2). Antenna: Length about 0.41 of head; shaft uniform up to middle and then gradually narrowed distally; uni- formly moderately pigmented, without spicules; all hairs single; hair 1 at about 0.5 from base. Thorax: Apices of long hairs attenuate; integument without spicules; stel- late hairs with few branches. Prothorax: hair 0(5, 4-6), 1(3), 2(1), 3(3, 3-5), 4(3, 2- 4), 5(3, 3-4), 6(1), 7(3, 2-3), 8(5, 5-8), 9(1, 1-2), 10(1), 11(2, 1-2), 12(1), 14(14, 10- 18). Mesothorax: hair 1(3, 3-4), 2(1), 3(1), 4(1), 5(1), 6(5, 4-5), 7(1), 8(5, 5-6), 9(5, 5-7), 10(1), 111, 1-2), 12(1), 13(7, 3-8), 14(5, 3-5). Metathorax: hair 1(4, 3-5), 2(1), 3(3, 3-5), 4(3, 3-5), 5(5, 3-6), 6(1), 7(5, 5-7), 8(6, 4-7), 9(4, 3-5), 10(1), 11(1, 1-2), 12(1), 13(4, 3-6). Abdomen: Stellate hairs with fewer branches than in fulvithorax. Segment I: hair 1(5, 4-5), 2(6, 3-6), 3(1), 4(4, 2-8), 5(6, 5-7), 6(4, 3-4), 7(1), 9(3, 3- 5), 10(1), 11(6, 5-8), 13(2, 2-5). Segment II: hair 0(1), 1(3, 2-4), 2(6, 5-8), 3(1), 4(3, 2-5), 5(5, 4-6), 6(3), 7(2, 1-3), 8(1), 9(4, 3-5), 10(1), 11(1), 12(1), 133, 2-5). Seg- ment III: hair 0(1), 1(2, 1-4), 2(7, 4-7), 3(1), 4(1), 5(5, 3-5), 6(2), 7(4, 2-5), 8(1), 9(6, 4-7), 101), 11(1), 12(1), 13(@2, 2-3), 14(1). Segment IV: hair 0(1), 1(2), 2(6, 5- 8), 3(1), 4€1), 5(4, 4-6), 6(2), 7(3, 3-5), 8(1), 9(7, 4-7), 10(1), 11(1), 1201), 13Q, 2- 3), 14(1). Segment V: hair 0(1), 1(2, 1-2), 2(7, 5-8), 3(1), 4(2, 1-2), 5(5, 3-5), 6(2), 7(3, 3-4), 8(1), 9(6, 4-7), 10(1), 11(1), 12(1), 13@, 1-3), 14(1). Segment VI: hair O(1), 1(2), 2(6, 6-7), 3(1), 4(@2, 1-2), 5(4, 3-5), 6(1, 1-2), 773, 3-5), 8(1), 9(4, 3-5), 10(1), 11(1), 12(1), 13(2, 2-4), 14(1). Segment VII: hair 0(1), 1(2), 2(6, 6-7), 3(1), 4(1), 5(3, 2-4), 6(2, 1-4), 7(1), 8(3, 2-5), 9(3, 3-4), 10(1), 111), 12(1), 13(2, 2-5), 14(1). Segment VIIT: Comb scales usually 20(13-27), in 3 irregular rows, elongate, rounded apically and evenly fringed; hair 0(1), 1(3, 2-3), 2(1), 3(5, 4-6), 4(1), 5(4, 3- 4), 14(1). Siphon: Index about 2.2-2.3; darkly pigmented and imbricate. Pecten ex- tending to about 0.4; usually 15 teeth (12-15), smaller at base and gradually becom- ing larger distally, all with 1-3 basal denticles. Hair 1(3, 2-3), 2(1), 3-5 not visible, 6(1), 7(1), 8(@2, 1-3), 9(1). Anal Segment: Saddle darkly pigmented and uniformly imbricate. Gills moderately long, narrow and subequal. Hair 1(2), 2(4, 3-5), 3(1), 4a(3, 3-4), 4b(2, 2-3), 4c(2, 2-3), 4d(2, 2-3), 4e(2, 2-3), 4£(2). SYSTEMATICS. In the present interpretation, septemstriatus appears to be re- stricted to areas of rain forest, chiefly in the lowlands, in Nicaragua, Panama and Co- lombia. In the past it has been confused with arborealis and aureolineatus but it ap- pears to be more closely related to fulvithorax on the basis of male genitalia and im- mature stages. This species has a more marked sexual dimorphism in adult ornamentation than any other member of the group, the male approaching the condition in both sexes of fulvithorax and the female having a striking arrangement of golden and dark bronzy scales in lines on the mesonotum. In the area of overlap of the 2 forms in Colombia, fulvithorax and septemstriatus are very strongly differentiated and there is no indi- cation of intergradation; for this reason septemstriatus is considered here to be a dis- tinct species. There is considerable variation in the ornamentation of the adults of septemstria- Ze Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 tus but the available material is too scanty to determine whether this is entirely in- dividual, ecological or geographical. However, it appears that in Colombia the light scaling is more restricted than in Panama and Nicaragua and particularly that the sex- ual dimorphism is not as marked. The above descriptions are based on individually associated material from Panama. Although the adults of septemstriatus are usually readily separated from all other members of the group on the diagnostic characters in the key and descriptions, I have not been able to find any reliable characters to separate it in the pupal stage from fulvithorax. The larvae of the 2 species are also very similar and it is possible that some of the characters used to differentiate them are not reliable because only the nonhairy form of septemstriatus has been seen. BIONOMICS. The immature stages of septemstriatus have been found in treeholes and broken bamboo. In one instance they were associated with 3 species of Haema- gogus, Aedes (F.) terrens, 2 species of Culex and a species of Sabethes (Galindo, Car- penter and Trapido 1955). The adults are diurnal, primarily sylvan, and are found mainly at the forest floor and not in the canopy (Trapido, Galindo and Carpenter 1955; Trapido and Galindo 1957). Other published reports on the bionomics of sep- temstriatus (Kumm and Novis 1938; Causey and Santos 1949; Floch and Fauran 1954; Belem Virus Lab 1964) do not apply to this species and should probably be attributed to arborealis. DISTRIBUTION (fig.2). Lowland to moderate elevations in Colombia (Upper Ori- noco drainage), Costa Rica (Pacific side), Nicaragua (Atlantic side) and Panama (At- lantic and Pacific sides). Material examined: 151 specimens; 44 6, 57 °, 42 pupae, 8 larvae; 12 individual rearings (9 pupal, 3 larval). COLOMBIA. Meta: Restrepo, 12 July 1935, W.H.W. Komp, 2 6 [USNM], 2 ? (KO 209A-9), 1° (KO 209A-11) [UCLA]; 22 July 1935, W.H.W. Komp, 9 6, 3 , 24; July 1935, 1 6 genitalia, 1 L; 22 Aug 1935, W.H.W. Komp, 1 6 [USNM], 1 ? (KO 206A-1), 4 6, 8 2 (KO 206A-7), 1 6 (KO 206A-13) [UCLA]; 2 May 1942, W.H.W. Komp, 2 2; 11 [USNM]. Villavicencio, 16 July 1943, W.H.W. Komp, | 9; 25 Oct 1943, M. Bates, 3 9; 1944, M. Bates, 1 p, 1 1 [USNM]. COSTA RICA. Puntarenas: Esquinas, elev. 300 ft, 6 June-30 Nov 1950, 1 A (Trapido, Galindo and Carpenter 1955:530). NICARAGUA. Bluefields, W.F. Thornton, 2 9? topotypes [USNM]. PANAMA. Bocas del Toro: Almirante, Bocas-Chiriqui Road, 2 May 1963, A. Quinonez (PA 301), 3 Ipd (301-106,109,112), 5 pd (301-102,105 108,110,111), 4 p? (301-101,103,104,109), 29 P. Chiriquisito, Trail Caldera, 5 Nov 1965, P. Orjuela (PA 1004), 5 9 [UCLA]. Canal Zone: Em- pire, 11 Feb 1926, J.B. Shropshire, 1 ¢. Fort Randolph, 27 July 1926, D.P. Curry, 1 6. France Field, 20 Feb 1926, J.B. Shropshire, 6 6, 19 9. Mojinga Swamp, 1 A (Blanton and Peyton 1958: 32). Paja (=Nueva Emperador), 13 May 1941, G.B. Fairchild, 5 d, 1 2. Summit, 30 May 1938, W.H.W. Komp, 1 ?; 7 May 1943, W.H.W. Komp, 1 6; 15 May 1945, W.H.W. Komp, 1 6; 1 ? [USNM] . Chiriqui: Chorcha, elev. 300 ft, 12 June-11 Dec 1950, 1 A (Trapido, Galindo and Car- penter 1955:530). David, C.S. Ludlow, 1 @ (Dyar 1923:148) [Army Medical Museum]. Cocle: Tucue, elev. 1000 ft, 12 June-14 Dec 1950, 14 A (Trapido, Galindo and Carpenter 1955:530). Colon: Candelaria, elev. 200 ft, F4-27 Aug 1950, 5 A (Trapido, Galindo and Carpenter 1955:530). Panama: Arraijan, elev. 600-800 ft, 5 June-29 Dec 1950, 2 A. Campana, elev. 2000 ft, 14 June-6 Dec 1950, 12 A. Campanita Arriba, elev. 600 ft, 24 Aug-4 Dec 1950, 14 A. La Victoria, elev. © 400-1200 ft, 7 Feb-28 Dec 1950, 1 A (Trapido, Galindo and Carpenter 1955:530). Pacora area, 10 2 (Galindo, Trapido and Carpenter 1950:548,551). San Jose Island: 26 July 1944, W.H.W. Komp, 1 9 [USNM]. 3. Aedes (Howardina) arborealis Bonne-Wepster & Bonne Figs.2,8,14,15 Berlin: Subgenus Howardina Pe 1920.Aedes arborealis Bonne-Wepster and Bonne, 1920:178. TYPE: Lectotype 6 (3893) with as- sociated larval and pupal skins (M 53) and genitalia slide (M 41), both slides marked BB 655, Dam, Surinam, Jan 1919, J. Bonne-Wepster and C. Bonne [ITH; selection of Belkin 1968]. Aedes (Howardina) arborealis of Bonne and Bonne-Wepster (1925:377-380); Dyar (1928:23 1-232); Edwards (1932:155); Lane (1936b:186; 1939:99); Floch and Abonnenc (1944:41; 1947:10); Lane (1953:708-710); Horsfall (1955:471); Stone, Knight and Starcke (1959:173); Floch and Fauran (1954:48); Fauran (1961:29); Belkin, Schick and Heinemann (1965:63); Forattini (1965:400); Stone (1967:208). Aedes septemstriatus of Kumm and Novis (1938:502,507,510-511); Causey and Santos (1949: 478); Floch and Fauran (1947:10; 1954: 48); Fauran (1961: 29); Belem Virus Lab (1964:101, 103,109,112). FEMALE. Wing: 2.8 mm. Proboscis: 2.45 mm. Forefemur: 1.93 mm. Abdomen: about 2.63 mm. In most respects intermediate between fulvithorax and septemstria- tus but differing from both in the following. Head: Median line of golden scales very narrow, these scales linear; submedian dark patches distinct but smaller than in sep- temstriatus. Thorax (fig.8): Mesonotum predominantly with golden yellow scales an- teriorly; ornamentation superficially very similar to 9 of septemstriatus except that golden acrostichal line narrower, fossa predominantly with bronzy scales, and mar- ginal golden area interrupted in front as well as caudad of scutal angle; caudal part of mesonotum with dark scaling more extensive than golden; upper supraalar patch of dark scales conspicuous. Scutellum with a conspicuous median line of golden scales. Scales of apn, ppn, and ppl golden, remaining pleural scales silvery; apn with numerous dense broad scales; ppn patch restricted, without broad scales. Abdomen: Median basal tergal patches as in septemstriatus on VI,VII but apparently small and yellowish on IV,V. MALE. Wing: 2.8 mm. Proboscis: 2.62 mm. Forefemur: 1.75 mm. Mesonotal or- namentation essentially as in the female but golden scales more numerous; acrostichal golden line broad; paraacrostichal dark-scaled line very narrow; outer dorsocentral dark line evident only posteriorly; fossa with small patch of dark bronzy scales; lower part of ppn with a few broad scales. Head: Without distinct dark patches, with only a few dark scales. Palpus subequal to proboscis in length, with numerous pale scales on lower basal surfaces of segments 3 to 5. MALE GENITALIA (fig.14). As figured; diagnostic characters as in the key. In general similar to fulvithorax except for the following. Ninth tergite lobe indistinct, with 6,7 strong bristles. Claspette with subapical seta subequal to apical but not markedly thickened; other setae much longer than in fulvithorax, apical seta extend- ing beyond middle of sternomesal margin of sidepiece. Clasper with about 5 setae in distal half. PUPA (fig.14). Abdomen: 3.24 mm. Trumpet: 0.44 mm; index 3.3-3.4. Paddle: 0.65 mm. Only nonhairy form known. Readily differentiated from fulvithorax and septemstriatus by fewer branches in hairs 1-II, 9-VII, VIII and broader paddle. Chae- totaxy as given below; hairs moderately to darkly pigmented. Cephalothorax: Meso- notum between trumpets moderately pigmented, darker laterad; metanotum moder- ately pigmented; trumpets darkly pigmented and strongly contrasting. Hair 1(1), 2(1), 3(1), 4(1, 1-2), 5C1, 1-2), 6(1, 1-2), 7(1, 1-2), 8(2, 2-3), 9(1), 10(2), 11(1), 12(1). Ab- domen: Tergite I moderately pigmented except for usual light membranous areas; other segments moderately pigmented and integument imbricate; pigmentation pro- gressively lighter caudad. Segment I: hair 1(5-7 main branches, subequal in length and each with many secondary branches), 2(1), 3(1), 4(2, 2-4), 5(1), 6(1), 7(1, 1-2), 24 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 9(1). Segment II: hair 0(1), 1(2, 1-4), 21, 1-2), 3(1), 4¢4, 1-4), 5(1), 6(1), 71, 1-2), 9(1). Segment III: hair 0(1), 1(1), 21), 31, 1-2), 4C1, 1-3), 51), 6(1), 7@, 1-4), 8@, 1-2), 9(1), 101, 1-2), 111, 1-2), 14(1). Segment IV: hair 0(1), 1(1), 2(1), 3@, 2-3), 41, 1-3), 5(1), 6(1), 71, 1-2), 8C1, 1-2), 9(1), 10€1), 11€1), 141). Segment V: hair O(1), 101), 201), 301), 4(@, 1-3), 3(1),, 61), 7G, 1-2), 8(@2, 122), 901), 10d), 110), 14(1). Segment VI: hair 0(1), 1(€1), 2(1), 31), 4€1), 51), 6(1), 7(1), 8C1, 1-2), 9(1), 101), 11(1), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(1), 4€1), 5C1), 6(1), 7Q1), 8(1), 9(2, 1-2), 10(1), 111), 141). Segment VUIT: hair 0(1), 4(1), 9(3), 14(1). Pad- dle: Ovoid; lightly pigmented, midrib dark; apex slightly produced; hair 1 single; male genital lobe extending to 0.6 of paddle, female genital lobe to 0.34. LARVA (fig.15). Only nonhairy form known. Readily differentiated from fulvi- thorax and septemstriatus by the following. Head hairs 6,11-C usually double; stel- late hairs with heavier spikes; comb scales usually more numerous (27-39), in 4,5 rows, siphon index about 2.0; 10-11 pecten teeth without basal denticles, the distal teeth greatly elongate; anal gills shorter than saddle, broad and subequal. SYSTEMATICS. As indicated in the diagnosis above, arborealis is somewhat inter- mediate between fulvithorax and septemstriatus in the ornamentation of the adults. Its larva and pupa are quite distinct from these species but may be similar to the un- known immature stages of aureolineatus which appears to be closely related to ar- borealis. In the present interpretation, arborealis has a more restricted distribution than for- merly reported, being confined to the Guianas and the lower Amazon. Former re- ports of arborealis from Colombia pertain to septemstriatus and all the specimens from the lower Amazon identified as septemstriatus in USNM appear to be arborealis. The above descriptions are based on the type series from Surinam. Specimens from Guyana and French Guiana agree in all respects with the Surinam material. The adults from the lower Amazon are usually considerably darker, with more extensive dark bronzy scaling on the mesonotum, but there is a great deal of variation in this character and some specimens are indistinguishable from the material from French Guiana. I consider all these populations as members of a single species. A single specimen of aureolineatus from Para without specified locality, suggests that this species and arborealis may overlap in the lower Amazon but there is no in- dication of intergradation of the 2 forms in the material at hand from Belem and up- river. | BIONOMICS. The immature stages of arborealis have been found in treeholes on- ly. There are no published reports on the adult bionomics of arborealis but the rec- ords of septemstriatus taken on human bait (Belem Virus Lab 1964) and during the wet (Oct-Mar) and dry seasons (Causey and Santos 1949) undoubtedly refer to ar- borealis in the present sense. DISTRIBUTION (fig.2). Confined to low elevations in Guyana, Surinam, French Guiana and Brazil (lower Amazon drainage). Material examined: 80 specimens; 8 6, 57 2, 6 pupae, 9 larvae; 7 individual rearings (3 larval, 4 incomplete). BRAZIL. Para: Belem, July 1931, R.C. Shannon, 1 @ [USNM]; IAN Forest, 413 ?; IAN and Utinga forests, 18 9; Utinga Forest, 107 ? (Belem Lab Report 1964:101,103,109,112). Curralinho, 1 Apr 1936, 1 9; Apr 1936, 1 ?; 26 July 1936, 1 3, 1 9; 1936, 1 L; 2 9 (K 325,341); Mar 1938, 5 9 [USNM]. Ilha de Marajo, 1 9 (Kumm and Novis 1938:507). Santarem, Belterra, Aug 1938, 1 9; Oct 1938, 1 9 [USNM]. FRENCH GUIANA. Baduel (Floch and Abonnenc 1944:4; 1947:10). Cayenne, Apr 1943, 1 9 [USNM] ; June 1944, E. Abonnenc, 32 9. Cabassou, 4 Feb 1965, T.H.G. Aitken, R. Martinez and A. Guerra (FG 46), 1 ¢ [UCLA]. Iracoubo (Fauran 1961:29). Placer Isnard, 17 June 1940, E. Abonnenc, 2 9 [UCLA]. Rorota, 8 May 1941, 2 9, 3 lp (577,578) [USNM]. Berlin: Subgenus Howardina 25 GUYANA. Mazaruni, 25 June 1936, W.H.W. Komp, 2 2? [USNM]; 6 July 1936, Leslie, 1 6, 2 9 (KO 208A-7); 6 July, 1 6 (KO 208A-8); 1 6 (KO 208A-10) [UCLA]. SURINAM. Aloesoebanja, 17 Jan 1956, E. van der Kuyp, 1 L. Dam, 1 Ip, “‘cotype’”? [USNM, No.22712]; 1 lp? (39-09, “‘cotype’’), 1 lp (39-01); 1 ¢ (38-98, “‘cotype”) [UCLA]. 4. Aedes (Howardina) aureolineatus Berlin, n.sp. Figs.2,8,58 . TYPES: Holotype ¢, Piraja (Bahia), Brazil, May 1931, Servico Febre Amarela, M.E.S., Brasil [USNM]. Allotype ¢ (660121-10) with genitalia slide, same data as holotype [USNM]. Paratypes: 2 2, same data as holotype [USNM, UCLA]. FEMALE. Wing: 3.6 mm. Proboscis: 2.7 mm. Forefemur: 2.32 mm. Abdomen: about 3.33 mm. Head: Median line of golden scales very narrow; these scales broader than in fulvithorax and curved; submedian dark patch very large, extending to occi- put; a small elongate lateral dark line at level of middle of apn. Palpus about 0.19 of proboscis. Torus light yellowish brown, darkened on mesal surface. Thorax (fig.8): Mesonotal integument brown; scaling predominantly dark bronzy; a conspicuous acrostichal golden line from anterior promontory, bifurcating on prescutellar space and reaching scutellum; a golden outer dorsocentral line arising caudad of scutal an- gle, broad and curved outward anteriorly, narrow posteriorly and reaching scutellum; lateral margin golden scaled, broadly to level of middle of ppn, narrowly to scutal an- gle, broadly caudad of scutal angle, separated by dark scales from a very large golden patch in and above supraalars; a few scattered golden scales laterad of acrostichal line especially in front. Scutellum with a conspicuous median line of narrow golden scales. Paratergite as in fulvithorax. Scales of apn numerous, light golden, moderately broad and curved; ppn patch extensive, light golden, scales short and curved; remain- ing pleural scale patches silvery to very light golden. Abdomen: Median basal tergal light patches silvery on VI,VII, light brown on V, usually absent on other segments. MALE. Wing: 3.06 mm. Proboscis: 3.1 mm. Forefemur: 2.16 mm. Allotype lack- ing proboscis, 1 antenna, palpi and all legs on right side; description from Salvador specimen. Essentially similar to female except that scattered golden scales more nu- merous and lateral marginal light line wider on mesonotum. Palpus subequal to pro- boscis in length. MALE GENITALIA (fig.58). As figured; diagnostic characters as in the key. Es- sentially as in arborealis except that lobe of tergite IX more prominent and several setae of the claspette distinctly thickened. IMMATURE STAGES. Unknown. SYSTEMATICS. The adults of aureolineatus are strikingly different from all other members of the group except sp. 5 (Iquitos form) in the reduction of the golden scaling on the central anterior part of the mesonotum to a narrow acrostichal line. This form appears to be a distinct species probably related to arborealis, but this can- not be definitely determined until the immature stages are found. In the small series of aureolineatus very little individual variation is evident. The Specimens from Para and Rio de Janeiro are not significantly different from the type material from Bahia. BIONOMICS. Nothing is known of the breeding sites and adult bionomics of au- reolineatus but it is likely that they are similar to other members of this group. DISTRIBUTION (fig.2). Confined to low elevations in Brazil along the coast from Para to Rio de Janeiro. 26 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 BRAZIL. Bahia: Piraja, May 1933, 1 d, 3 9, type series [USNM, UCLA]. Salvador, 27 Apr 1936, E. Silva, 1 6 [USNM]. Para: Apr 1930, N.C. Davis, 1 ° [USNM] . Rio de Janeiro: Apr 1933, 1 9 [USNM]. 5. Aedes (Howardina) sp., Iquitos form FEMALE. Wing: 3.33 mm. Proboscis: 2.71 mm. Forefemur: 2.34 mm. Abdomen: broken off. Extremely similar to aureolineatus and differing apparently chiefly in the following. Mesonotal ornamentation darker; acrostichal golden line narrower; dorso- central golden line represented only by a rounded spot at about 0.5 of mesonotum; fewer scattered golden scales; apn scales more numerous and broader, more distinctly silvery; ppn scale patch more extensive, with some broad scales below. MALE and IMMATURE STAGES. Unknown. DISCUSSION. The single specimen from Iquitos, Peru, Mar-Apr 1931, R.C. Shan- non [USNM] lacks the abdomen. It may represent a distinct species related to arbor- ealis and aureolineatus or a local population of one of these. This form shows the greatest reduction in the golden scaling on the anterior part of the mesonotum in the Fulvithorax Group. WALKERI SECTION FEMALES. Usually small to medium sized species, wing 2.1-3.5 mm. Head: Sub- median dark patch broad; lateral patch smaller, bordered posteriorly by light scales. Erect scales varied. Torus brown to dark brown, with a few small black scales mesal- ly. Palpus with or without apical light scales. Thorax: Mesonotum usually with dis- tinct linear light pattern (except in some grabhami), light scales all golden or combi- nation of golden and silvery. Acrostichal line absent but a small light patch present at level of anterior promontory; inner and outer dorsocentral lines incomplete, inner not reaching scutellum, outer usually not reaching anterior margin; lateral marginal line broken at scutal angle, usually narrow (very broad in Walkeri Group); median prescutellar line present. Paratergite bare. Acrostichal bristles usually absent on disc, present in Busckii Group. Pleural scales silvery (fig.17); ppn and psp without scales; ssp scale patch present or absent; mep with a small, restricted patch at upper half be- low bristles only. Legs: Midfemur and hindfemur without distinct distal light spot on anterior surface. Foretibia sometimes with light scales distally. Tarsal light markings not restricted to segment 1. Wing: Veins entirely dark scaled. Abdomen: Sternites H- IV without dark scales. MALES. Larger claws of foreleg and midleg with both submedian and basal teeth present (in Busckii Group basal tooth absent). MALE GENITALIA. Lobes of tergite IX indistinct, bearing a few strong setae. Sidepieces separated from each other at base by a distance greater than diameter of aedeagus at that level; tergomesal and sternomesal margins without any indication of lobes. Claspette represented by a small roughly triangular or fingerlike lobe bearing 1, rarely 2, apical setae; not connected to its mate. PUPAE. Cephalothorax: Hair 5-C usually shorter than distance between its base and that of trumpet; either subequal to or longer than hair 4-C; if longer, varied in branching. Abdomen: Hair 5-IV,V usually single (double or triple in Aurites and Busckii Groups), terminating before or extending beyond apex of tergite following; Berlin: Subgenus Howardina | 8 | 2-IV-VI usually distad of 0.75 of its tergite; 9-III-VI usually caudolaterad of hair 6 (except in walkeri); 9-VII moderately to strongly developed, either close to or re- moved considerable distance cephalad of hair 6. LARVAE. Stellate hairs developed or not. Antenna: Short; with or without scat- tered spicules; hair 1-A single. Thorax: Integument glabrous. Tubercles of hairs 5-7-P distinctly joined; hair 9-P simple, usually single or double; hair 13-P absent except in Ioliota Group. Abdomen: Integument glabrous except in walkeri and members of Ioliota Group. Hairs 2,5,9-VII usually stellate; hair 8-VII cephalad or cephalodorsad of 9-VII. Segment VIIT: Hairs 1,5-VIII usually strong, with a few branches but not stellate. Comb scales usually in a single row (up to 2-4 irregular rows in Busckii Group); individual scale usually blunt apically. Siphon: Index 2.0-8.5. Integument smooth, imbricate or spiculate. Acus distinct except in Walkeri and Aurites Groups. Pecten teeth with or without basal denticles. Anal Segment: Saddle with distinct marginal spicules. Hair 1-X short to moderate, with a few branches; hair 3-X single, 2-X multiple. Ventral brush with 5 or 6 pairs of hairs arising from a boss with or without distinct grid bars. DISCUSSION. The Walkeri Section is the most heterogeneous of the 3 sections in the subgenus, particularly in the immature stages. However, it is probably a mono- phyletic lineage characterized by 3 unique features of adult ornamentation: parater- gite and ppn both without scales and prescutellar area with a median light-scaled line. Four rather distinct phylads within the section are suggested by differences in the mesonotal pattern of the adults: (1) Aurites Group, with a rather broad linear golden pattern very reminiscent of the Sexlineatus Section, (2) Walkeri Group, with a very broad continuous lateral silvery patch and apparently relatively broad golden dorso- central lines, (3) the very similar Ioliota and Albonotatus Groups, with a very narrow silvery lateral line, very faint golden dorsocentral lines and a silvery prescutellar line, and (4) Busckii Group, intermediate between the Walkeri Group and the Ioliota- Albonotatus Groups in the width of the lateral silvery line which is frequently broken. The Busckii Group is unique in the subgenus in the presence of acrostichal bristles on the disc of the mesonotum. The immature stages are much more diverse than in the other sections, each of the above-mentioned phylads possessing a type quite as distinct as those characterizing the Sexlineatus and Fulvithorax Sections. The Ioliota Group is also clearly separated in the larval and pupal stages from the Albonotatus Group. The breeding sites in the Walkeri Section encompass the entire range used by the subgenus, extending to habi- tats not utilized by the other sections. The Walkeri Section is almost entirely confined to islands in the Caribbean area. Only 2 small encroachments on present continental areas are known: (1) in the Yu- catan Peninsula by the Ioliota Group and (2) in northern Venezuela and Trinidad by the Ioliota Group and northern Venezuela only by the Walkeri Group. The groups show primarily allopatric complementary distributions. The only exceptions known are in (1) the Walkeri and Aurites Groups in Jamaica, (2) possibly Albonotatus and Busckii Groups in Guadeloupe, and (3) Ioliota and Walkeri Groups in northern Vene- zuela. Except in the Aurites Group, members of a group always have allopatric dis- tributions (fig.3). There are 2 surprising gaps in the distribution of the Walkeri Section. Although in- tensive collections have been made in Puerto Rico, Virgin Islands and Leeward Is- lands, no species of Howardina has yet been recorded from this area. Nor has the subgenus been recorded from Cuba. However, it is possible that the record of Haema- gogus sp. from Cuba by Montchadsky and Garcia (1966:46-47) is based on a mis- 28 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 identification of an undescribed species of Howardina. The diversity of types in the Walkeri Section is undoubtedly a reflection of typical insular speciation through isolation of small populations. As indicated above, it ap- pears that only one original stock was probably involved, although there is a sugges- tion of a possible recent intrusion of the Sexlineatus Section in the Aurites Group (see). However, the present pattern could not have been attained through a simple splitting of the original stock among the island masses as they exist today. The reten- tion of such primitive features as acrostichal bristles in the adults of busckii, hair 13- P in the larvae of the Ioliota Group, 6 pairs of hairs in the ventral brush in all species except busckii and a patch of comb scales in the larvae of some populations of busckii, is indicative of very early separation of the Walkeri Section from the other sections rather than its recent derivation from one of them. That the present pattern represents speciation and dispersal on island masses no longer in existence in the Caribbean area is strongly supported by the relict distributions of the Walkeri and Ioliota Groups. KEYS TO GROUPS AND SPECIES ADULTS 3 All mesonotal leht scaling golden (Aurntes Group)... 2°. we At least lateral marginal lime silvery-or white <0 2. 2 7) cosue Teeio nw a Aurites Group 2(1). Pleuron with a large silvery patch continuous from lower pra across posterior stp toitslower margin. . . - oc 6) aurives Pleuron with 3 small discrete, separate silvery parches. one each on lower Pra, Upoct. se and {ower 870 (6 Oe A ee 3(2). Antealar area (above paratergite) with golden scales, rarely 1-3 bronzy scales present; mesonotal light lines usually distinct and broad, sometimes almost completely merging in anterior two-thirds of mesonotum . .9. grabhami Antealar area with numerous dark bronzy scales above narrow posterior lat- eral mesonotal golden line; mesonotal light lines narrow, discrete, never Mere ie ee Oe IO imeeamalis, 1 ic siene 4(1). Lateral marginal line broadened to form a conspicuous lateral silvery patch; ssp without scales (Walkeri Group) . Lateral marginal line very narrow, not extending o on fossa: - ssp with patch of ls yes atin cen ee Walkeri Group 5(4). Inner dorsocentral line continuous, moderately broad. . >. . . 6. walkeri Inner dorsocentral line dotted, only faintly indicated . . . . . 7. argyrites Busckii Group 6(4). Median prescutellar line golden, faint; palpus entirely dark in both Sexes; acrostichal bristles presenton disc @ 2) 0 a eee a 12 busekil 7(6). 8(7). (7). oul S(2). 4(3). =), Berlin: Subgenus Howardina 2 Median prescutellar line silvery, conspicuous; silvery scales on apex of female palpus and base of some segments of male palpus; acrostichal bristles ab- Sent OHCne a ee ee Hindtarsal segment 5 with ae scales (Ioliota Group) eg ee ae Hindtarsal segment 5 with black scales only (Albonotatus Group). re Ioliota Group Hindtarsal segment 5 entirely white-scaled . . . tues 6G 2. toloda Hindtarsal segment 5 with some apical and posterior dark scales . ws comnclensis Albonotatus Group Hindtarsal segment 3 white-scaled in barely more than basal 0.5 . ; Mee ee a) eg Hindtarsal segment 3 white-scaled in at least basal0.7. . . 14. albonotatus MALE GENITALIA (7. argyrites not definitely known) Tergal surface of sidepiece with very dense vestiture of long bristles, some as long as’sidepiece . . >. 5 a. LD. Daehamensis Tergal surface of sidepiece with sparse vestiture of short or moderate bristles, none more thay.0.5’of sidepiece length 27 ee? Gk ee Clasper very a including spiniform about as peice as sidepiece . 14. albonotatus Clasper short or r moderate, including spiniform not more than 0.75 of side- eieceleneih.. . -. Caete cromp ik Ged (OILS peak ig eine ee Claspette with an apical fingerlike cylindrical process more than twice as long as wide and bearing apical bristles; body of claspette with short subapical Drisiies: 2 ae. 4 Claspette without distinct apical fingerlike cylindrical process, apical bristle borne on a short broad conical process; body of claspette with several strong subapical bristles borne on similar but smaller tubercles . . . .5 Apex of aedeagus deeply emarginate in dorsal aspect producing lateral lobes and/or distal half of aedeagus distinctly broader than proximal half; basal tergal bristles of sidepiece strong, subequal in length and development to tergomesal marginal bristles. . . . pu Soe EO ser Aedeagus not emarginate apically in dorsal aspect and roughly of uniform width; basal tergal bristles of sidepiece weak, much shorter and weaker than tergomesal marginal bristles 6. aurites, 9. grabhami; | 0. inaequalis; : a stenei Clasper more than 0.5 of sidepiece length, without distinct subapical setae . 6. walkeri Clasper less than 0. 5 of sidepiece length. with distinct subapical eetae <0 30 6(5). 2(1). 3(2). 4(3). a¢3). 6(2). 7(6). 8(6). 9(8). Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Clasper gradually tapered todistalO.75 ..... . .13. cozumelensis Clasper swollen on base, abruptly tapered at about middle So wh pedoge PUPAE (7. argyrites unknown) Walkeri Group Paddle deeply emarsinate athar!] .°. . 6. . . . 6. walkeri Paddle usually smoothly rounded on apex, at ctost shallowly emarginate at PR a a ei as ee ee ds ee ee Hair 10-C usually at least 10-branched (Aurites ee Sy se ak a peace Hair 10-C usually less than 5-branched. . .. . ee en eek Aurites Group Pigmentation very light and uniform; metanotum pene scape pigmen- ted ore. Pigmentation moderate and t uneven; metanotum with light < area in the middle er ro a ee Hair 2-¥ idistinctly laterad Of Nair 3. 4 ea ee ses hae Ge oe © UTES Flaw 2-Vi miesad or at level of Neir3. . 6 Om es BR Bona Median tergal darkening of abdominal segment III indistinct. ie SC, inaequalis Median tergal darkénine of abdominal segment i sharply marked | 11. stenei Hair 5 at least 2.0 of hair 4 (loliota Group). . . ae aoe Hair 5-C subequal to hair 4 (Albonotatus and Busckii Groups). Sree Ioliota Group Hair 1-II strongly dendritic and with many branches; paddle ovoid, length about 1.5 greatest width . . i G2. toon Hair 1-II with 7-12 simple branches: ‘paddle rounded, length about 1.2 great- OS Wiltd ue Beak Begs err Gay ah ane, hy eyes Albonotatus and Busckii Groups Hairs 7,10-C both Single; paddle elongate, length more than 1.5 greatest width . . 14. albonotatus Hairs 7 10-C usually at least double: paddle length moderate, usually less than 1. 3 greatest Widths so ucan he ee Pinna very long, more than 0.33 of total trumpet length; paddle slightly emarginate at hair 1; hair 6-III],IV strong, usually at least double . . . L535. bahamensis Pinna moderate, about 0. a of total trumpet length: paddle evenly rounded on apex; hair 6-III AV Weak Wives Pe ea ee, Base 2(1). erg 4(2). 5(4). 6(5). At). 8(7). 9(8). Berlin: Subgenus Howardina 31 LARVAE (7. argyrites unknown) Walkeri Group Abdominal integument densely covered with large dark-branched spicules . oc Bia Gm Me Te ee el ey ee: 6. walkeri Abdominal injeeument Slanraus at A a a ee es ee ee Hair 13-P present (loliota ack ae WT ee RESET OO NO COE ini Sarna NACE RE Hair 13-P' absent ... 3 fMea Gk eet lea ige Ugly eas ees ae eae Ioliota Group Siphon slightly swollen in middle, index about 2.5; comb scales blunt or rounded apically... . ped £2 Aptliota Siphon slightly evenly tapered from base, index. about a OQ; comb scales Ponte APIO ee I I aa ions Busckii Group Ventral brush with 5 pairs of hairs (comb scales sometimes in 2 to 4 rows; pecten teeth sometimes extending well beyond hair 1-S). . . 16. busckii entrar Dish) Will G paws OL ais” Ce ee eee Head hair 6-C usually single or double; pecten teeth short and with basal den- ticles; siphon index 3.0 or less (Albonotatus Group) . . .6 Head hair 6-C multiple, usually more than 5-branched; pecten teeth long, slender and without basal denticles; siphon index more than 4.0 (Aurites eC) petunia ee ee at Albonotatus Group Anal gills subequal; siphonal hair 1-S adjacent to last pecten tooth . . . . 14. albonotatus Anal ills markedly unequal siphonal hair 1. -S well beyond last pecten tooth ool Eh ae ERR cs £5, bahamensis Aurites Group Hair 2-VII large, stellate, 4-7-branched. . ...... . .. .. Jd. stenei PROT ee i IR OR ie iar dyna Wie ecw aga a age eee ee Siphon index less than 5.5; anal gills markedly unequal, the small distinctly shorter than saddle... |... . . . 10. inaequalis Siphon index 6.0 or more; anal gills subequal and distinctly longer than sad- 1 Nill idle Coe ihdib inline ree Rue) a Bi) Lene Head hair 5-C usually single; 7-C very short, less than 0.25 of antenna, and usually triple . . ay? grabhami Head hair 5-C a at least triple; +e about 0. 5 of antenna and usually 6- DYANENOG 464. eae gc ee Na ie ae ee ee ee ae 32 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Walkeri Group FEMALES. Head: Narrow decumbent scales white. Palpus with apical silvery scales (broken in argyrites). Thorax: Inner dorsocentral line golden, continuous or dotted; median prescutellar line broad, white or predominantly white; outer dorso- central line faint, narrow, golden; lateral marginal line broadened to form a conspic- uous lateral silvery patch from anterior angle to wing base. Acrostichal bristles not developed on disc: Scutellum predominantly dark scaled, light scales restricted to median lobe. Ssp without scales; upper stp patch separate, large. Legs: Foretarsus completely dark; midtarsus with incomplete or complete basal white ring on segment 1; hindtarsus with moderate basal white rings on segments 1-3 in walkeri, at least on 1-2 in argyrites (rest broken). Abdomen: Basomedian tergal light markings creamy, present on proximal segments. MALE (walkeri only). Palpus with basomedian. white scales on segments 4,5. Larg- er claw of foreleg and midleg with both submedian and basal teeth; smaller claw with basal tooth only. MALE GENITALIA (walkeri only). Caspette with short broad conical tubercle bearing 1 apical bristle. Clasper about 0.6 of sidepiece; minute spicules basally; sub- apical hairs absent. Aedeagus subquadrate, shorter than proctiger, without apical ~ lobes. PUPA (walkeri only). Cephalothorax: Hair 5-C usually at least triple, moderately long, subequal to distance from its alveolus to trumpet base and at least 2.0 as long as hair 4; hair 7-C usually 3-branched; hair 10-C with 2-7 branches. Abdomen: All large hairs except 1-II,IJI, 9-VII,VHI usually single or double; hair 2-II laterad of or in line with hair 3; hair 5-[V,V usually extending to apex of tergite following; hair 9- II-VI usually at level or slightly cephalad of hair 6; 9-VII subequal to hair 6-VIJ; hair 4-VIII single. Paddle: Short; apex distinctly emarginate; marginal spicules distinct. LARVA (walkeri only). Head: Hair 1-C always branched, usually triple; 4-C un- usually long, multiple; 5-C single, long and stout; 6-C usually 4-branched; hairs 11, 12,14-C all stellate, 11-C with unequal branches; 15-C double, long, exceeding apex of mental plate. Antenna: Shaft with scattered, short, sharp spicules; hair 1-A single. Thorax: Hair 13-P absent. Hairs 0,1,3,4,8,14-P, 1,13,14-M, 1,3,4,5,8,13-T all strong- ly stellate, with numerous branches, very darkly pigmented; hairs 1-M,T far mesad of ~ hairs 2,3. Abdomen: Integument with numerous very dark, large branched spicules; hairs 2 and 9 removed cephalad of hair 8. Hairs 1,2,5,9,13-I-VI, 11-I, 7-II-VI all strongly stellate as on thorax; hair 8-VII cephalodorsad of 9-VII. Segment VUIIT: Comb scales in a single row; individual scale long, pointed and fringed to apex. Si- phon: Index less than 3.0; integument with rows of short spicules; acus absent, no indication of basal indentation. Pecten extending almost to apex; teeth long, proxi- mal ones simple, distal branched apically. Anal Segment: Saddle body with numer- ous strong spicules; caudal margin with long fringed spines; hair 1-X multiple, near lower end of caudal margin. Ventral brush with 6 pairs of hairs on very strongly scle- rotized boss without distinct grid bars. DISCUSSION. The diagnosis of the group is based almost entirely on walkeri, the type species of the subgenus and a common species endemic to Jamaica. A. argyrites, known only by the imperfect female holotype from northern Venezuela and possi- bly a male genitalia slide, is extremely similar to walkeri in adult ornamentation and is undoubtedly a member of this group. The Walkeri Group possesses many features unique in the section, particularly in the immature stages which are readily recognized from all other species in the sub- Berlin: Subgenus Howardina 33 genus. However, the mesonotal pattern of the adults suggests a greater affinity with the Aurites Group than with any other group. As indicated under the latter, this is interpreted here as an indication of introgression with the Sexlineatus Section. The widely disjunct distribution of the 2 members of the Walkeri Group parallels that of the 2 members of the Ioliota Group and is similarly interpreted as a relict pat- tern of a group originally with a wide distribution in the central Caribbean area (fig. B). The group may be primarily a breeder in bromeliads as is the case with walkeri but since the latter sometimes uses treeholes it is possible that these may have been the original breeding sites, perhaps still retained by argyrites. 6. Aedes (Howardina) walkeri (Theobald) Figs.3,7,8,16-18 1901.Culex walkeri Theobald, 1901:424. TYPE: Holotype ? (45/110), Jamaica, date and collector not specified [BM]. Aedes (Howardina) walkeri of Bonne and Bonne-Wepster (1925:368); Dyar (1928:235); Edwards (1932:156); Lane (1939:101; 1953:725-726); Horsfall (1955:471); Stone, Knight and Starcke (1959:174); Belkin, Schick and Heinemann (1965:31); Forattini (1965S: 399 ,400); Porter (1967: 38). Aedes walkeri of Dyar and Knab (1906b:189,192); Howard, Dyar and Knab (1912:70; 1917:849- 852); Picado (1913:325); Dyar (1918:73,80); Johnson (1919:424); Gowdey (1926:74); Thomp- son (1947:78); Hill and Hill (1948:46). Howardina walkeri of Theobald (1903a:287-289); Dyar (1905a:49; 1905b:23; 1905c:27-28); Theo- bald and Grabham (1905:20-22); Theobald (1905:9; 1907:215-216; 1910:220); Coquillett (1906b:26); Grabham (1906:170-171); Surcouf and Gonzalez-Rincones (1911:139-140). Culex walkeri of Blanchard (1905:416). Haemagogus walkeri of Dyar and Knab (1906a:166). FEMALE (fig.16). Wing: 2.62 mm. Proboscis: 2.1 mm. Forefemur: 1.75 mm. Ab- domen: about 2.45 mm. As described for group, with the following apparently diag- nostic features. Head: Median stripe on vertex broad, silvery; light scales on sides and venter creamy. Thorax (figs.8,16): Scales on anterior promontory golden. Inner dor- socentral line distinct, continuous and moderately broad; median prescutellar line usually silvery, sometimes with a few golden scales, rarely predominantly golden; outer dorsocentral line golden; lateral marginal line very broad, silvery. Antealar area above paratergite with silvery scales. Silvery scales on median scutellar lobe narrow; lateral lobes with narrow dark scales. Pleural scales as in the group. Legs: Forefemur and midfemur predominantly dark anteriorly, a few basal scales creamy, proximal 0.5 of posterior surface broadly creamy, extending distally as a light line, rest dark; hindfemur mostly creamy except for lateral distal dark scales. Midtarsal segment 1 with only a few white scales at base. Haltere: Stem pale, knob mostly dark, fringe scales creamy white. Abdomen: Basomedian tergal creamy patches distinct on {I-VI a few pale scales on II. Sternites creamy on II-IV, rest of segments with basal creamy and apical dark scales. MALE (fig.16). Wing: 2.28 mm. Proboscis: 1.93 mm. Forefemur: 1.04 mm. Essen- tially similar to female in color. MALE GENITALIA (fig.17). As figured; diagnostic characters as in the key. Ninth tergite lobe indistinct, with 3,4 strong setae. Sidepiece about 2.5 times median width. 34 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Clasper with minute spicules basally; spiniform about 0.2 of clasper. PUPA (fig.17). Abdomen: 2.7 mm. Trumpet: 0.38 mm; index about 4.5-5.0. Pad- dle: 0.46 mm. Chaetotaxy as figured; hairs moderately to darkly pigmented. Diagnos- tic characters as in the key. Readily separated from all other species of the subgenus by the presence of minute denticles on abdomen and by the distinctly emarginate paddle. Cephalothorax: Mesonotum moderately pigmented between trumpets, slight- ly deeper laterally; trumpets contrasting. Hair 1(1), 2(2, 2-3), 3(1, 1-2), 4(3, 2-5), 5(5, 3-6), 6(1, 1-2), 7(3, 2-4), 8(3, 1-5), 9(1), 1005, 2-7), 1101, 1-2), 12(3, 2-5). Ab- domen: Tergite I moderately to strongly pigmented except for usual membranous areas; remaining segments moderately to strongly pigmented, progressively lighter caudad; integument with minute denticles. Segment I: hair 1(8-12 primary branches, each with many secondary branches), 2(1, 1-2), 3(1), 4(3, 2-5), 5(4, 3-6), 6(1, 1-2), 7(2, 1-3), 9(1, 1-2). Segment II: hair 0(1), 1(8,.5-17), 2(1), 31), 4(@2, 2-4), 5@, 1-3), 6(1, 1-2), 7(3, 1-4), 81, present in some), 9(1). Segment III: hair 0(1), 1(4, 2-9), 2€1, 1-2), 3(1), 4@, 1-3), 5(2, 1-3), 6(1), 7(2, 2-5), 8(3, 2-5), 9(1), 10(1, 1-2), 11(1), 14(1). Segment IV: hair 0(1), 1(3, 2-4), 2(1, 1-3), 3(3, 2-6), 4@, 1-2), 5(2, 2-3), 6(1, 1-2), 7(2, 1-4), 8(2, 2-3), 9(1), 101, 1-3), 11€1), 14(1). Segment V: hair 0(1), 1(2, 1-3), 2(1, 1-3), 3(1, 1-3), 41, 1-5), 5@, 2-4), 6(1, 1-2), 7(2, 2-5), 8(2, 2-4), 9(1), 101, 1- 2), 111), 14(1). Segment VI: hair 0(1), 1(1, 1-3), 2(1, 1-3), 31, 1-2), 4(2, 1-4), 5(3, 2-4), 6(1, 1-2), 7(2, 1-3), 8(2, 2-3), 9(1), 10(1), 11€1), 14(1). Segment VII: hair 0(1), 1(1, 1-3), 21, 1-3), 3(2, 1-3), 4(1, 1-2), 5(@2, 1-4), 6(5, 3-6), 7(1, 1-2), 8(2, 2-6), 9(6, 6-9), 10(1), 11(1, 1-4), 14(1). Segment VIII: hair 0(1), 4(1), 9(12, 11-16), 14(1, 1-2), Paddle: Uniformly lightly pigmented, midrib slightly darker; dorsal surface with fine denticles, external margin serrated; apex deeply emarginate; hair 1-P slender. Male genital lobe extending to 0.5 of paddle, female genital lobe to 0.41. LARVA (fig.18). Head: 0.7 mm. Siphon: 0.43 mm. Anal Saddle: 0.22 mm. Chae- totaxy as figured; hairs darkly pigmented. Diagnostic characters as in the key. Readi- ly differentiated from all other species of the subgenus by: (1) stellate hairs with nu- merous branches, (2) dendritic spicules on abdomen and (3) pecten teeth extending almost to apex of siphon. Head: Width subequal to length; integument smooth and strongly pigmented. Mental plate with 12 distinct teeth on each side of strong medi- an tooth, often a smaller detached tooth at base. Hair 0(1), 1(3, 2-3), 3(1), 4(6, 5-8), 5(1), 6(4, 2-5), 7(4, 3-5), 8(1), 9(2), 10(€1), 11(14, 9-17; stellate, branches of 3 lengths), 12(10, 8-17), 13(1), 14(15, 11-24; stellate), 15(2, 1-3). Antenna: Length about 0.3 of head; uniformly strongly pigmented; with scattered spicules; all hairs usually single; hair 1-A about 0.62 from base. Thorax: Integument glabrous; in life thorax appears bluish green. Apex of long hairs attenuate; stellate hairs with numer- ous branches; long and stellate hairs weakly barbed. Prothorax: hair 0(32, 30-46), 1(32, 27-38), 2(1), 3(26, 24-36), 4(36, 30-42), 5(4, 3-5), 6(1, 1-2), 7(3, 2-4), 8(36, 30-38), 9(2, 1-4), 10(1), 11(4, 3-5), 12(1), 14(48, 44-54). Mesothorax: hair 1(36, 25- 54), 2(1), 31), 4C1), 5(1), 6(8, 5-8), 7(1), 8(8, 7-10), 9(10, 7-14), 10(1), 11(3, 2-4), 12(1), 13(22, 17-27), 14(23, 19-29). Metathorax: hair 1(54, 48-63), 2(1), 3(22, 19- 24), 4(28, 28-34), 5(25, 22-33), 6(1), 7(8, 7-10), 8(19, 16-23), 9(8, 6-8), 10(1, 1-2), 11(3, 2-4), 12(1), 13(42, 38-43). Abdomen: Integument densely packed with large dendritically branched spicules, more sparsely on segment I; stellate hairs as on tho- rax. Segment I: hair 1(53, 51-71), 2(40, 40-53), 3(1), 4(2, 1-2), 5(29, 28-41), 6(3, 3-4), 71, 1-2), 9(14, 9-15), 10(1), 11(26, 24-30), 13(35, 29-38). Segment II: hair O(1), 1(60, 52-65), 2(46, 36-46), 3(1), 4(2, 1-3), 5(32, 25-36), 6(4, 3-4), 7(18, 14- 22), 8(1), 9(20, 18-22), 10(1), 11(1), 12(1), 13(36, 33-44). Segment III: hair 0(1), 1(64, 56-66), 2(41, 35-45), 3(1), 4(1), 5(34, 30-40), 6(3, 2-3), 7(20, 20-24), 8(1), Berlin: Subgenus Howardina 35 9(21, 21-26), 10(1), 11(1), 12(1), 13(36, 32-44), 14(1). Segment IV: hair 0(1), 1(60, 54-62), 2(42, 36-46), 3(1), 4(1, 1-2), 5(37, 30-42), 6(2, 2-3), 7(20, 18-22), 8(1), 9(24, 22-27), 10(1), 11(1), 12(1), 13(40, 25-41), 14(1). Segment V: hair 0(1), 1(60, 47-61), 2(37, 37-42), 3(1), 4(3, 1-3), 5(36, 30-41), 6(@2, 2-3), 7(20, 18-25), 8(1), 9(26, 25-31), 10(1), 11(1), 12(1), 13(36, 30-42), 14(1, 1-2). Segment VI: hair 0(1), 1(56, 44-60), 2(41, 36-41), 3(1), 4(2, 1-2), 5(30, 30-42), 6(2, 1-2), 7(22, 18-24), 8(1), 9(28, 25-29), 10(1), 111), 1201), 1334, 28-38), 14(1). Segment VII: hair 0(1), 1(50, 42-50), 2(36, 30-40), 3(1), 4(1), 5(28, 28-39), 6(20, 16-26), 7(1), 8(24, 22- 28), 9(26, 26-32), 10(1), 1101), 12(1), 13(22, 20-28), 14(2). Segment VIIT: Comb scales in a single row of 12(10-14); individual scales long, pointed apically, fringed to the tip. Hair 0(1), 1(48, 46-54), 2(1), 3(8, 8-10), 4(1), 5(28, 26-31), 14@, 1-2). Si- phon: Index about 2.8-2.9; darkly pigmented; integument uniformly pilose. Pecten extending beyond hair 1-S, almost to apex; 20(16, 22) teeth, progressively longer dis- tad, individual teeth long, without basal denticles, distal teeth broad, with apical den- © ticles. Hair 1-S about 0.67 from base. Hair 1(3, 2-4), 2(1), 3-5 not visible, 6(1), 7(1), 8(1), 9(1). Anal Segment: Saddle strongly pigmented, with darker basal ring; integu- mentary spicules progressively larger distally; caudal margin with long spines fringed laterally. Gills narrow, moderately long, ventral pair slightly shorter than dorsal. Hair 1(8, 6-10; strong, at caudolateral angle), 2(12, 10-14), 3(1), 4a(7, 6-8), 4b(7, 6-7), 4c(6, 5-8), 4d(6, 6-7), 4e(6, 6-7), 4f(S, 4-7). EGG. Length about 0.62 mm, width about 0.14 mm; exochorion transparent and endochorion dark and with polygonal sculpturing. SYSTEMATICS. A. walkeri, the type species of Howardina, is among the most clearly marked species in the subgenus. In the adults, only the closely related argyri- tes, known by a single female from Venezuela, could be confused with walkeri. The immature stages are also very strikingly differentiated but it would not be surprising if those of argyrites, when found, were to conform to the same general type in view of the resemblance of the females. Little variation is apparent in the extensive material of walkeri, either in the adults or the immature stages. Only the median prescutellar line exhibits some noticeable variation, ranging from entirely silvery in the majority of specimens to predominant- ly golden in a few. The dorsocentral lines are always clearly marked and never dotted - aS in argyrites. BIONOMICS. The breeding sites of walkeri are usually bromeliad axils but a few collections of immatures have been made in treeholes (JA 336,390), broken and cut bamboo (JA 711,778), heliconia axils (JA 748) and a crabhole (JA 369). Both terres- trial and arboreal bromeliads are used by walkeri. This species has been found associ- ated with all members of the Aurites Group and appears to have a much wider alti- tudinal and geographical range than any other species of Howardina on Jamaica. The larvae of walkeri are readily recognized in the field by their very dark and “hairy” bodies. Females of walkeri commonly bite humans under the shade of bromeliad laden trees. I found a maximum peak of activity in mid morning (0900) and a lesser one in late afternoon (1700) as shown in fig.7. However, Hill and Hill (1948:46) state that walkeri prefers to feed at dusk in the open, and bites at any time of the day in forest shade. DISTRIBUTION (fig.3). Endemic to the island of Jamaica where it is widespread at elevations from 100 ft to 4500 ft. Material examined: 2964 specimens; 244 4, 1139 2, 699 pupae, 882 larvae; 516 individual rearings (237 pupal, 150 larval, 129 incomplete). 36 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 JAMAICA. Clarendon: Honeycomb Rock, elev. 800 ft, 8 Aug 1965, W.A. Page (JA 237), 1 Ip (237-27) [UCLA]. Mocho Road, 10 Sept 1965, J.N. Belkin, 1 9 (JA 353) [UCLA]. Slippery Fall, elev. 800 ft, 8 Aug 1965, W.A. Page, 1 L (JA 239) [UCLA]. Hanover: Copse, elev. 500 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson (JA 668), 1 pd (668-101), 1 p? (668-100), 6 L, 21 [UCLA]. Lethe, elev. 500 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson, 4 p, 1 L, 5 1(JA 655), 1 lp? (667- 10), 5 pd (667-100-104), 5 IP (667-11-15), 2 9,2 P, 4 p, 34 L, 41(JA 667) [UCLA]. Manchester: Hope Farm, elev. 1000-1100 ft, 10 Sept 1965, W.A. Page, 1 p? (354-102), 1 IP (354-13), 55 9 (JA 356); 23 Oct 1966, O.G.W. Berlin and D.C. Watson, 31 2 (JA 634A), 2 Ipd (635-11,16), 2 Ip? (635- 10,12), 4 pd (635-100-103), 1 p? (635-104), 8 IP (635-13-15,17-21), 1 P, 21 (JA 635); 2 Nov 1966, O.G.W. Berlin, 12 2 (JA 663A), 779 2 (JA 684) [UCLA]. Mandeville, elev. 900 ft, 23 Oct 1966, O.G.W. Berlin and D.C. Watson (JA 639), 4 lp? (639-10,14,22,25), 8 pd (639-101,104,107- 111,114), 7 p? (639-100-103,105,106,112,113), 12 IP (639-11-13,15-21,23,24), 9 6, 4 9, 18 P, 22 p, 62 L, 24 1; 24 Oct 1966, O.G.W. Berlin (JA 640), 1 IP (640-10), 1 L [UCLA]. Melrose Hills, elev. 900 ft, 8 Aug 1965, W.A. Page (JA 244), 1 lpd (244-43), 1 p? (244-41A), 1 IP (244-40), 7 P [UCLA]. Porus, elev. 900-1000 ft, 8 Aug 1965, W.A. Page (JA 243), 2 Ipd (243-22,29), 3 lp? (243-21 ,26,27), 1 pd (243-102), 2 p? (243-101,103), 2 IP (243-19,20), 1 P, 15 L; 23 Oct 1966, O.G.W. Berlin and D.C. Watson, 1 IP (631-30), 2 Ip? (633-11,12), 1 p? (633-100), 5. IP (633-10,13- 16), 4 L (JA 633); 1 Nov 1966, O.G.W. Berlin (JA 657), 2 lpd (657-10,11), 1 pd (657-100), 2 p? (657-101,102), 2 IP (657-13,14), 2 P, 2 p, 1 L, 7 1; 2 Nov 1966, O.G.W. Berlin, 1 lpd (660-10), 1 pd (660-100), 1 P (JA 660), 1 Ip? (664-12), 4 pd (664-100,103-105), 2 p? (664-101,102), 2 IP (664-10,11), 1 P, 22 L(JA 664) [UCLA]. Williamsfield, elev. 1100 ft, 23 Oct 1966, O.G.W. Berlin and D.C. Watson (JA 637), 4 Ipd (637-10,11,13,17), 3 lp? (637-12,23,27), 10 pé (637-100-105, 108-110,112), 3 p? (637-106,107,111), 11 IP (637-14-16,18-22,24-26), 3 p, 25 L, 1 1 [UCLA]. Portland: Ecclesdown, elev. 1000 ft, 22 Aug 1967, J.N. Belkin and W.A. Page (JA 748), 1 p? (748- 94), 1 p [UCLA]. Fairy Hill, elev. 100 ft, 13 Nov 1966, O.G.W. Berlin and D.C. Watson, 20 L, 21 (JA 693), 4 Ipd (694-11,13,14,17), 9 lp? (694-10,12,15,18-23), 21 pd (694-100-103 ,106-108,112- 117,119-121,123,125,128-131), 13 p? (694-101 ,104,105 ,109-111,118,122,124,126,132,133), 1.1P (694-16), 4 6,1 9,2 P, 2 p, 34 L (JA 694), 19 L (JA 695) [UCLA]. Kemney-Sherwood Forest, elev. 750 ft, 13 Aug 1967, J.N. Belkin and W.A. Page (JA 707), 3 lpd (707-10-12), 2 pd (707-100, 102), 1 IP (707-13), 1 P, 1 L; 13 Aug 1967, W.A. Page, 2 Ipd (708-11,12), 1 lp? (708-15), 1 d(JA 708), 1 Ipd (709-10), 1 pd (709-101), 1 p, 1 L(JA 709), 1 L(JA 710), 1 lpd (716-10), 3 pd (716- 100,101,104), 1 P, 4 L (JA 716) [UCLA]. Portland Point, 4 Aug 1942, C.B. Philip, 1 @ [USNM]. St. Andrew and Kingston: Castleton, elev. 600 ft, 27 Oct 1966, O.G.W. Berlin and D.C. Watson (JA 645), 2 IP (645-10,11), 1 P, 5 L, 5 1 [UCLA]. Duhaney River, 6 Nov 1965, W.A. Page (JA 399), 1 IP (399-13) [UCLA]. Ferry Quarry, 13 Sept 1965, W.A. Page, 1 d (JA 369) [UCLA]. Hardwar Gap, elev. 3800 ft, 29 Oct 1966,0.G.W. Berlin (JA 647), 2 lpd (647-10,11), 1 lp? (647- 12), 1 pd (647-100), 1 p? (647-101), 1P, 1 p, 1 L, 11 [UCLA]. Hermitage Dam, 1934, 6 9? [USNM]. Hermitage Dam Road, 22 Aug 1965, W.A. Page (JA 282), 1 Ipd (282-36), 1 Ip? (282-33), 1 IP (282- 31); 13 Aug 1965, W.A. Page, 1 IP (294-44), 1 IP (304-21); 1 Sept 1965, W.A. Page, 1 Ipd (319-22), 1 Ip? (319-23), 1 pd (319-104), 31 L(JA 319), 1 lp? (322-23), 3 IP (322-24,31,32), 1 9 (JA 322), 3 L (JA 323), 2 p? (324-102,103), 6 L (JA 324); 7 Sept 1965, J.N. Belkin (JA 333), 1 lp? (333- 21), 1 pd (333-100), 2 p? (333-101 ,102), 1 IP (333-20), 2 L; 29 Oct 1965, W.A. Page, 1 9 (JA 390), 1 P (JA 391), 1 Ipd (392-12), 1 Ip? (392-11), 6 pd (892-102,104,106,109,111,113), 4 p? (392- 100,101,105,114), 1 IP (392-10), 2 3, 2 9,2 p, 16 L(JA 392), 1 lpd (393-12); 15 Aug 1967, J.N. Belkin and W.A. Page, 1 Ip? (723-30), 1 pd (723-103), 1 p? (723-102), 2 L(JA 723), 2 Ipd (724- 12,14), 3 Ip? (724-10,11,15), 3 pd (724-102-104), 1 IP (724-13), 1 P, 2 p,3 L(JA 724), 2 Ipd (725-15,16), 1 Ip? (725-14), 5 pd (725-90,100A,105A,106,108A), 3 p? (725-92,107,112), 3 3, 3 2,1 P,3 p (JA 725); 1 Sept 1967, W.A. Page and J.N. Belkin (JA 765), 1 IP (765-10), 2 L; 13 July 1967, W.A. Page (JA 843), 3 lpd (843-12,13,15), 3 Ip? (843-10,11,14), 1 pd (843-101), 2 p? (843-100,102); 25 July 1967, W.A. Page, 1 L (JA 854), 1 Ipd (855-11), 2 lp? (855-1012), 3 IP (855-13-15), 1 P, 6 L (JA 855) [UCLA]. Kingston, 1 Apr 1906, M. Grabham, 1 6, 2 9 [USNM]. Mavis Bank, M. Grabham, 1 6 [USNM]. Mt. Salus Road, elev. 1500 ft, 11 Sept 1967, W.A. Page, 1 pd (777-100), 2 IP (777-10,11), 1 Ipd (779-10), 1 pd (779-104), 2 p? (779-101,105), 1 2, 1 P, 2 L (JA 779); 24 July 1967, W.A. Page (JA 848), 1 IP (848-10), 1 P, 1 L, 11 [UCLA]. Newcastle, Berlin: Subgenus Howardina 37 elev. 2500-4000 ft, 5 Sept 1964, W.A. Page (JA 124), 1 Ipd (124-15), 3 lp? (124-10,12,14), 1 IP (124-11), 2 L; 29 Oct 1966, O.G.W. Berlin (JA 648), 1 lpd (648-16), 1 lp? (648-19), 2 pd (648- 100,101), 9 IP (648-10-15,17,18,20), 1 P, 9 L, 13 1; track to Catherines Peak, elev. 4000-4300 ft, 8 Nov 1966, O.G.W. Berlin, 1 P, 1 p, 1 L, 11 (JA 678), 9 2 (JA 679), 2 p? (680-100,101), 1 9,2 P, 1 p, 2 L(JA 680), 1 p,2L,51(JA 681), 1 P, 16 L(JA 682), 1 P, 1 L, 11(JA 683) [UCLA]. Stony Hill, elev. 1100 ft, 18 July 1964, L. Paton (JA 85), 3 pd (85-101,103,104), 8 p? (85-100,102,105- 108,110,111), 1 P, 3 L;1 Sept 1965, W.A. Page (JA 318), 2 Ipd (318-32,35), 1 Ip? (318-38), 5 pd (318-33,34,110,111,113), 6 p? (318-102,104,106-109), 1 IP (318-36), 1 6,8 P, 1 p,5 L [UCLA]; 1934, 3 2 [USNM]. Uester Road, elev. 4500 ft, 8 Sept 1965, J.N. Belkin, 1 9 (JA 336), 5 lpd (337- 10,14,15,18,24), 2 lp? (337-11,16), 2 pd (337-22,23), 1 p? (337-27), 1 IP (337-17), 5 6,3 9, 1 P, 4 L (JA 337) [UCLA]. Waterworks track, elev. 1100 ft, 8 Sept 1965, J.N. Belkin (JA 335), 2 Ip (335-10,20), 3 lp? (335-11,22,30), 1 pd (335-100), 1 IP (335-25), 8 6, 2 2, 2 p,6 L [UCLA]. Woodcutters Gap, elev. 4500 ft, 3 Sept 1964, W.A. Page (JA 122), 1 lp? (122-11), 2 d(JA 122), 1 Ipd (123-10), 1 Ip? (123-11), 1 pd (123-100) [UCLA]. St. Ann: Moneague, elev. 500-1750 ft, 9 Sept 1965, D.C. Watson, 2 p? (606-100,101), 4 IP (606-10-13), 4 L (JA 606), 1 2? (JA 607); 17 Aug 1967, J.N. Belkin and W.A. Page, 2 L (JA 733); 31 Aug 1967, W.A. Page, 2 L (JA 762); 2.0 miS of Moneague near Unity, elev. 1750 ft, 28 Aug 1967, W.A. Page (JA 758), 1 Ipd (758-22), 1 Ip? (758- 21), 1 pé (758-102), 1 IP (758- 20): 29.0 mi post on Moneague Road, elev. 1750 ft, 30 Aug 1967, W.A. Page, 5 Ipd (763-10,11,13,15,17), 1 lp? (763-16), 1 pd (763-91), 4 p? (763-93,104,110 113), 36,19%,3p,4L; TIGA 763). 2 Ip? (764-11,20), 2 L(JA 764); Moneague-Claremont Road, elev. 750-1250 ft, 17 Sept 1965, W.A. Page, 1 6 (374-100), 1 Ipd (379-12), 2 lp? (379-10,14), 1 IP (379- 11), 4 LIA 379), 11(JA 381); 5 Aug 1967, W.A. Page, 1 p (JA 767) [UCLA]. St. James: Monte- go Bay, elev. 500 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson (JA 669), 1 p? (669-103), 1 IP (669-20), 2 L [UCLA]; Jan 1940, W.H.W. Komp, 21 [USNM]. Reading, elev. 200 ft, 6 Nov 1966, O.G.W. Berlin, 1 1(JA 670), 11(JA 671), 2 9 (JA 672) [UCLA]. St. Mary: Broadgate, elev. 500 ft, 25 Aug 1965, D.C. Watson, 1 L (JA 257); 27 Oct 1966, O.G.W. Berlin and D.C. Watson (JA 643), 2 Ipd (643-10,14), 4 Ip? (643-11-13,15), 1 pd (643-109), 2 p? (643-110,111), 1 IP (643-16), 2 p, 1 L, 81 [UCLA]. Castleton, elev. 600 ft, 27 Oct 1966, O.G.W. Berlin and D.C. Watson (JA 644), 1 Ip? (644-20), 1 p? (644-110), 6 P,4 L [UCLA]. St. Thomas: Barrett’s Gap, 14 Nov 1946, G.A. Thompson, 1 ? [IJ]. Norris, elev. 500 ft, 30 Oct 1966, O.G.W. Berlin and D.C. Watson (JA 653), 1 Ip? (653-21), 1 p? (653- 100), 2 IP (653 -20,22), 1 P, 4 p, 1 L, 1 1 [UCLA]. Pleasant Hill, elev. 250 ft, 14 June 1931, Kisliuk, 7 2 [USNM]. Port Morant Bay, 16 Aug 1967, J.N. Belkin and W.A. Page, 2 6,1L(JA 729) [UCLA]. Scotland Gate, elev. 500 ft, 30 Oct 1966, O.GW. Berlin and D.C. Watson (JA 654), 5 pd (654-100,102,103,107,109), 5 p? (101,104-106,108), 10 IP (654-10-19), 1 ¢,10P, 101 [UCLA]. Windsor Forest, elev. 750 ft, 30 Oct 1966, O.G.W. Berlin and D.C. Watson, 1 Ipd (650- 10), 1 pd (650-113), 1 p? (650- oe 4 IP (650-12- 15), 1 P,1L,21(QA 650), 4 Ipd (651-10,11,14,15), 5 lp? (651-12,17,19-21), 7 pd (651-100-102,105- 108), 5 pe (65 1-103,104,109, 115 116), 3 IP (651-13,16,18), 4 P, 1 p,6 L,11€JA 651) [UCLA] . Westmoreland: Whitehouse, elev. 100 ft, 11 Sept 1965, W.A. Page, L& (JA 359), 3 2 (JA 360) [UCLA] . Parish not specified: Sandy Folly, Feb 1945,G A. Thompson, 1 L; Jan 1940, W.H.W. Komp, 1 6; 1940, W.H.W. Komp, 1 9; Jan 1946, G.A. Thompson, a a 1940, 1 ?;M. Grabham, 1 6, 3 2, 1p, 51; R.B. Hill, 3 2; G.A. Thompson, 1 2 [USNM]; W.A. Page, 1 IP (771-11), 1 Ipé (772. 20), 1 lp? (772-21), 3 L. a (JA 772), 2 lpé (787-12,14), 2 Ip? (787-11,13), 3 pd (787-108,109,111), 7 p? (787-100-102, 104, 105,107,112), 1 IP (787-10), 1 P, 1 p, 10 L(JA 787) [UCLA]. 7. Aedes (Howardina) argyrites Dyar and Nunez Tovar Figs.3,8 1927.Aedes (Howardina) argyrites Dyar and Nunez Tovar, 1927:190. TYPE: Holotype ?, Vene- zuela, M. Nunez Tovar [USNM]. Aedes (Howardina) argyrites of Dyar (1928:230); Edwards (1932:155); Lane (1939:99); Anduze (1941:14); Lane (1953:708); Horsfall (1955:471); Stone and Knight (1956:226); Stone, Knight 38 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 and Starcke (1959:173); Belkin, Schick and Heinemann (1965:74); Cova Garcia, Sutil and Rausseo (1966a:58; 1966b:141,333). FEMALE. Apparently distinct from walkeri in the following features (fig.8). Sides and venter of head mottled with black and white scales; inner dorsocentral mesonotal line faint, narrow, dotted; midtarsal segment 1 with complete basal ring. The single known specimen has the apex of the proboscis, palpi, antennae and segments 3-5 of hindtarsi broken; hindtarsal segments 1 and 2 have basal white rings. MALE, PUPA, LARVA unknown. SYSTEMATICS. The pattern of ornamentation of the single known adult of argy- rites is so similar to that of walkeri that I have no hesitation in assigning argyrites to the Walkeri Group. The 2 forms are apparently differentiated primarily by the diag- nostic characters given above but since the holotype of argyrites is damaged it is pos- sible that additional differences may be present, especially in tarsal markings. A slide of male genitalia [USNM] from an unspecified locality in Venezuela (P.J. Anduze) probably represents another specimen of argyrites, as identified by W.H.W. Komp. There appear to be some differences between these and the male genitalia of walkeri but without additional material and the absence of the whole adult I hesitate to de- scribe them at this time. , The exact type locality of argyrites is not known but the most likely place is somewhere in the Cordillera de la Costa in the state of Aragua, the origin of the bulk of the material collected by Nunez Tovar. Failure to find additional material of argy- rites in this relatively well-known area suggests that argyrites may be a rare relict form. BIONOMICS. The holotype was presumably obtained in a biting-landing collec- tion. Immature stages of argyrites should be looked for in bromeliad axils, treeholes and broken bamboo, the known breeding sites of the closely related walkeri. Aurites Group FEMALES. Head: Narrow decumbent scales yellowish. Palpus with a few white scales on apex. Thorax: All light mesonotal lines golden, varied in development; lat- eral line sometimes moderately to rarely strongly broadened but not silvery. Acros- tichal bristles not developed on disc. Scutellum with linear golden scales on lateral lobes. Ssp with or without scales; upper stp patch separate or united. Legs: Foretar- sus completely dark; midtarsus with basal white markings on segments 1 or 1,2; hind- tarsus with basal white markings on segments 1,2 or 1-3. Abdomen: Basomedian ter- gal light markings when present restricted to distal segments. MALES. Palpus entirely dark. Larger claw of foreleg and midleg with small acute basal tooth and large submedian tooth; smaller claw with basal tooth only. MALE GENITALIA. Claspette with a distinct outstanding apical fingerlike cylin- drical process more than twice as long as wide and bearing a short apical bristle. Clasper about 0.5 of sidepiece; basal half with minute spicules; subapical hairs absent. Aedeagus subquadrate, slightly wider at middle, subequal to proctiger in length. PUPAE. Cephalothorax: Hair 5-C at least 4-branched, moderately long but usual- ly not reaching base of trumpet; hair 7-C with 3 or more branches; hair 10-C usually at least 10-branched. Abdomen: Hair 1-II-[V with many branches, 1-II with about as many branches as float hair (1-I); hair 2-II always mesad of hair 3; hair 5-IV,V usual- ly double and about 1.5 of tergite following; hair 9-II-VI usually at level or slightly Berlin: Subgenus Howardina 39 cephalad of hair 6; 9-VII subequal to hair 6-VII; hair 4-VIII double. Paddle: Moder- ate; practically unpigmented; apex rounded; marginal spicules indistinct. LARVAE. Head: Hair 1-C unbranched; 4-C short; 5-C single or branched, long; 6-C at least 6-branched, moderate; none of ventral hairs stellate; 14-C single; 15-C short, not extending cephalad of base of mental plate. Antenna: Shaft with a few scattered spicules; hair 1-A spikelike, single. Thorax: Hair 13-P absent. Only hairs 0, 1,3,14-P and 5-T stellate, with a few, weakly barbed branches; hair 1-T far mesad of 2,3-T. Abdomen: Integument glabrous; all hairs except 0,8,14 at about same trans- verse level. Only hairs 2,5-I-VI, 11-I, 7-9-II-VI stellate, with a few weakly barbed branches; hairs 2 and 9 stronger than others; 9-I small, single; hair 8-VII distinctly cephalodorsad of 9-VII. Segment VIII: Comb scales in a single row; individual scales long and fringed to apex. Siphon: Index 5.0-8.5; integument glabrous; acus absent but base of siphon with small lateral indentation. Pecten not reaching middle of si- phon; teeth long, without denticles. Anal Segment: Saddle body without distinct spicules except distally; caudal margin with row of long spines; hair 1-X always sin- gle and at lower end of caudal margin. Ventral brush with 6 pairs of hairs on a weak boss without grid bars. DISCUSSION. The clearly marked Aurites Group shows a phenomenon unique in the subgenus: a flock of 4 sibling species sympatric in one small area. To date the group has been recognized only in Jamaica, Were some of the species are quite com- mon (fig.3). There is a suggestion of some affinity of this group with the sympatric Walkeri Group in the general similarity in the mesonotal pattern of the adults. However, the immature stages of the 2 groups are very distinct. There is a much greater resem- blance in the mesonotal pattern of the adults with the Sexlineatus Section except for the prescutellar line, and the larvae and pupae also resemble the Sexlineatus Section more than any group in the Walkeri Section. Although this resemblance may be en- tirely fortuitous, 2 more plausible explanations are possible. First, the Aurites Group might be the primitive stock of the Walkeri Section derived from the Sexlineatus Sec- tion; this is very unlikely since the entire Sexlineatus Section lacks several important primitive characters represented in other groups of the Walkeri Section (acrostichal bristles of the adults and comb scales in a patch in the larva). Second, the view held here; the Aurites Group may have been formed through relatively recent introgres- sion between the Sexlineatus Section and the Walkeri Group. This interpretation would also account for the presence of a species flock in the Aurites Group, since such radiation usually takes place soon after a new phyletic line is established. It is also supported indirectly by the presence in Jamaica of several species in other gen- era that are either identical with or very closely related to Central American species, strongly suggesting a relatively recent connection between these areas. No specific differences in the male genitalia of members of the group have been found and the adults of inaequalis and stenei cannot be separated from each other. In other respects the 4 species are readily differentiated from one another as indi- cated in the keys. There appear to be some ecological barriers separating 2 of the species. A. inae- qualis breeds in treeholes, broken bamboo and rockholes, largely at low elevations, and never in bromeliad axils, which are utilized exclusively by the other 3 species. A. aurites breeds primarily at elevations above 3,500 feet and has never been found associated with the other bromeliad-breeding members of the group. A. grabhami and stenei occur primarily below 2,500 feet but grabhami has been collected at elevations up to 4,000 feet. These 2 species are usually found associated in collections but it is not known if they occupy the same axils under natural conditions. 40 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 8. Aedes (Howardina) aurites (Theobald) Figs.3 ,8,19,20 1907.Howardina aurites Theobald, 1907:216-218. TYPE: Lectotype ?, Newcastle (St. Andrew), Jamaica, elev. 4000 ft, July 1906, Col. Loscombe [BM; selection of Belkin, 1968]. Aedes (Howardina) aurites in part of Dyar (1928:236); Edwards (1932:155); Lane (1939:99; 1953: 720-722); Horsfall (1955:471). Aedes aurites of Howard, Dyar and Knab (1917:859-860); Dyar (1918:73); Johnson (1919:424); Gowdey (1926:73). Aedes aurites in part of Hill and Hill (1948:29,42). Howardina aurites of Theobald (1910:220); Surcouf and Gonzalez-Rincones (1911:139-140). Aedes (Howardina) inaequalis in part of Thompson (1956:199-200); Stone, Knight and Starcke (1959:174); Belkin, Schick and Heinemann (1965:31); Forattini (1965 :399,400). Aedes inaequalis in part of Stone and Knight (1956:219). FEMALE. Wing: 2.6 mm. Proboscis: 2.00 mm. Forefemur: 1.8 mm. Abdomen: about 2.7 mm. Head: Light decumbent scales golden on median stripe and short or- bital line, yellowish on sides and venter. Erect scales yellowish, a few lateral ones darker. Palpus about 0.16 of proboscis. Thorax (fig.8): Mesonotal dark scales dense; light lines golden, moderately broad, wider than dark lines; inner dorsocentral line reaching to prescutellar space, followed by moderately broad median prescutellar line; outer dorsocentral line complete, merging with lateral marginal line anteriorly; lateral marginal line long, extending from side of anterior promontory to wingbase, very broad from scutal angle to wingbase due to presence of golden scales on antealar area above paratergite. Median scutellar lobe with a row of narrow golden scales flanked on sides by broad dark ones; lateral lobes with a few narrow, golden and black scales. Apn with a distinct silvery patch; ssp bare; patches on base of pra, up- per and lower stp merged and continuous; rest of patches as in the section and group. Legs: Forefemur and midfemur predominantly dark anteriorly except for a few basal creamy scales, posterior surface and hindfemur predominantly creamy except for dis- tal dark areas. Midtarsus with few white scales on segment 1; hindtarsus with short white rings on 1,2, rest dark scaled. Haltere: Stem pale, knob predominantly dark, fringe scales whitish. Abdomen: Basomedian tergal light scaling inconspicuous. Ster- nites II-IV straw yellow, V predominantly yellow with few apical dark scales; rest of segments with basal creamy and distal dark scales. MALE. Wing: 2.34 mm. Proboscis: 2.16 mm. Forefemur: 1.71 mm. Essentially similar to female in coloration. MALE GENITALIA (fig.19). As figured; diagnostic characters as in the key. Ninth tergite lobe indistinct, with 2 strong bristles. Sidepiece length about 3.0 median width. Clasper about 0.6 of sidepiece, basal 0.3 with minute spicules; sly ae short, about 0.25 of clasper. PUPA (fig.19). Abdomen: 2.48 mm. Trumpet: 0.38 mm; index 6.0-6.2. Paddle: 0.7 mm. Readily separated from inaequalis and stenei by the absence of lightly pig- mented area on metanotum and hair 9-C single; from grabhami by hair 2-VI laterad of 3-VI. Cephalothorax: Mesonotum and metanotum uniformly lightly pigmented; trumpets contrasting. Hair 1(1), 2(2, 1-2), 3(1, 1-2), 4(5, 3-8), 5(5, 4-7), 6(1, 1-2), 7(4, 4-6), 8(1), 9(1), 10(10, 9-16), 11(1), 12(3, 2-6). Abdomen: Tergite I uniformly lightly pigmented except for usual membranous areas; rest of segments light, pigmen- tation progressively lighter caudad. Segment I: hair 1(12-18 primary branches, each with many secondary ones), 2(1), 3(1), 4(6, 3-8), 5(6, 2-6), 6(1, 1-2), 7(2, 2-3), 9(1), Berlin: Subgenus Howardina 41 10(1, present in some). Segment II: hair 0(1), 1(37-56 branches, variable in different specimens), 2(1), 3(1), 4(4, 2-4), 5(1), 6(1), 7(4, 3-4), 9(1), 10(1, present in some). Segment III: hair 0(1), 1(14, 6-24), 2(1), 3(1, 1-3), 4(4, 2-4), 5(1), 6(1), 7(3, 2-4), 8(3, 2-4), 9(1, 1-2), 10(2, 2-3), 11(1), 14(1). Segment IV: hair 0(1), 1(8, 5-16), 21, 1-2), 3(4, 3-6), 4(2, 1-4), 5(1, 1-3), 6(1), 7(3, 2-5), 8(2, 1-3), 9C1), 10@, 2-4), 11Q1), _ 14(1). Segment V: hair 0(1), 1(2, 1-7), 2(1, 1-2), 3(2, 2-3), 4(5, 3-8), 5(1), 6(1), 744, 2-6), 8(2, 1-3), 9(1), 10(€1, 1-2), 11(1), 14(1). Segment VI: hair 0(1), 1(1, 1-6), 2(1), 3(2, 1-3), 4(4, 2-6), 5(2, 1-5), 6(1, 1-3), 7(1), 8(3, 1-8), 91), 10(1), 1101), 14(). - Segment VII: hair 0(1), 1(1), 2(1), 31, 1-2), 4(2, 1-2), 5C1, 1-2), 6(2, 2-4), 7(1, 1-2), 8(3, 2-4), 9(3, 2-4), 10(1), 11(1), 14(1). Segment VIII: hair 0(1), 4(2), 9(10, 8-12), 14(1). Paddle: Almost without pigmentation; midrib faintly pigmented; external margin smooth. Male genital lobe extending to 0.42 of paddle, female genital lobe to O27: LARVA (fig.20). Head: 0.73 mm. Siphon: 0.97 mm. Anal Saddle: 0.24 mm. Read- ily separated from stenei by hair 2-VII small and single; from inaequalis by the longer subequal gills and at least 3-branched hair 5-C; from grabhami by the 3-branched hair 5-C. Chaetotaxy as given below, hairs moderately pigmented. Head: Width about 1.18 of length; pigmentation light, deeper around mouthparts and behind ocular area ventrally. Mental plate with a strong median tooth and 10 distinct teeth on either side. Hair 0(1), 1(1), 3(1), 4(4, 4-5), 5(3, 2-4), 6(10, 9-13), 7(6, 4-7), 8(1), 9(3, 2-3), 10(1), 11(3, 3-5), 12(1, 1-2), 13(1), 141, small), 15(3, 3-4). Antenna: Length about 0.44 of head, very lightly pigmented and with few scattered spicules. All hairs single; hair 1-A about 0.5 from base. Thorax: Long hairs attenuate apically and weakly barbed; stellate hairs with a few weakly barbed branches. Prothorax: hair O(11, 8- 14), 109, 9-11), 21), 313, 12-14), 41), 5Q, 2-3), 6(1), 7(1, 1-2), 8(8, 8-12), 9Q, 1- 3), 10(1), 11(4, 2-5), 12(1), 14(16, 11-19). Mesothorax: hair 1(5, 3-6), 2(1), 3(1), 4(1), 5(1), 6(5, 4-6), 7(1), 8(7, 6-8), 9(8, 6-9), 10(1), 11(2, 1-3), 121), 13(10, 6-16), 14(5, 5-9). Metathorax: hair 1(2, 1-4), 2(1), 3(4, 3-5), 4(7, 3-9), 5(9, 6-10), 6(1), 7(7, 6-9), 8(4, 3-5), 9(5, 4-7), 10(1), 11@2, 1-2), 121), 13(7, 4-7). Abdomen: Stellate hairs as on thorax. Segment I: hair 1(3, 2-3), 2(7, 6-8), 3(1), 4(4, 2-5), 5(7, 7-10), 6(3, 3-4), 7(1), 9C1, 1-2), 10(1), 11(7, 6-9), 13(5, 3-6). Segment II: hair 0(1), 1(@2, 1- 3), 2(7, 6-8), 3(1), 4(2, 2-4), 5(9, 7-10), 6(3), 7(3, 2-4), 8(1, 1-2), 9(6, 3-7), 10(1), 111), 12(1), 133, 3-5). Segment III: hair 0(1), 1(2, 2-3), 2(6, 5-8), 3(1), 4(@2, 2-3), 5(10, 8-10), 6(1, 1-2), 7(5, 4-7), 8(1), 9(6, 5-6), 10(1), 11(1), 12(1), 13(5, 3-6), 14(1). Segment IV: hair 0(1), 1(2, 2-3), 2(7, 6-7), 3(1), 4(2, 2-3), 5(10, 8-11), 6(1, 1-2), 7¢4, 4-8), 8(1, 1-2), 9(6, 5-7), 10(1), 11€1), 121), 13(5, 4-6), 14(1). Segment V: hair O(1), 1(2), 2(6, 5-7), 3(1), 4(2, 2-5), 5(9, 7-10), 6(1), 7(3, 3-5), 8C1, 1-2), 9(6, 5-6), 101), 11(€1), 121), 13(5, 4-7), 14(1). Segment VI: hair 0(1), 1(2), 265, 4-6), 3(1), 4(1), 5(6, 5-9), 6(1), 7(2, 2-3), 8(3, 1-3), 9(4, 4-6), 10(1), 11(1), 121), 13(7, 5-8), 14(1). Segment VII: hair 0(1), 1€1, 1-2), 2(1), 3(1), 41), 5(3, 2-4), 6(3, 2-3), 7(1), 8(6, 4-7), 9(3, 2-3), 10(1), 11(1), 12(1), 133, 2-5), 14(1). Segment VIIT: Comb scales in a single row, 18(16-19) in number; individual scale long and fringed uniformly to apex. Hair 0(1), 1(4, 3-5), 2(1), 3(6, 4-6), 4(1), 5(2, 2-3), 14(1). Siphon: Index about 6.0-6.5; moderately pigmented, with darker basal ring. Pecten extending to about 0.45, with 19(14-21) teeth; proximal teeth smooth, distal with a few apical spinules. Hair 1(2, 2-3), 2(1), 3-5 not visible, 6(1), 7(1), 8(2, 1-3), 9(1). Anal Segment: Saddle moderately pigmented; caudal margin with spines of varying sizes. Gills subequal, long, about 2.0 of saddle length. Hair 1(1), 2(5, 4-7), 3(1), 4a(4, 4-5), 4b(4), 4c(4, 3-5), 4d(4, 3-5), 4e(4, 3-4), 4f(3, 2-4). SYSTEMATICS. A. aurites is undoubtedly a distinct species erroneously synony- 42 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 mized by Thompson (1956:199) with inaequalis. It is readily differentiated in the adult stage from all the other members of the group, as pointed out by Theobald (1907:217) in the original description, by the large continuous silvery patch on ssp. There are other diagnostic features in the adults as indicated above and the immature stages are readily separated from the other members of the group. There is no striking variation in adult ornamentation in the small series of speci- mens examined and the diagnostic characters of the immature stages are quite con- stant. BIONOMICS. The immature stages of aurites have been found only in arboreal bromeliads, in my experience generally at elevations of more than 3,500 feet and never in association with grabhami or stenei, the other bromeliad-breeding members of the Aurites Group. It appears that this species occurs at lower elevations at least occasionally as Thompson reared several adults from larvae collected at Hope Farm [USNM] at an elevation of 1,000 feet in 1946, and 2 females (JA 702) were caught recently (1967) near Kingston at elevations of about 1,000-1,500 feet. Females have been collected biting humans during the day but no attempt has been made to study their activity cycle. From the available data, aurites appears to have a restricted geo- graphic range, having been collected only from the parishes of Manchester and St. Andrew. DISTRIBUTION (fig.3). Endemic to the island of Jamaica where it has been re- ported from the parishes of Manchester, St. Andrew and Kingston, primarily at ele- vations of 3400 to 5250 ft. Material examined: 145 specimens; 24 6, 33 9, 44 pupae, 44 larvae; 36 individual rearings (22 pupal, 7 larval, 7 incomplete). JAMAICA. Manchester: Hope Farm, 1 Jan 1946, G.A. Thompson, 2 Ipd (460101-24,26), 1 1d (460101-30), 1 lp? (460101-27), 1 lp (460101-15); 1 Oct 1946, G.A. Thompson, 1 p (461001-1) [USNM]. St. Andrew and Kingston: Cinchona Gardens, elev. 4000-5250 ft, 8 Sept 1965, J.N. Bel- kin, 1 d (JA 341); 24 Aug 1967, W.A. Page, 3 L (JA 752) [UCLA]. Ferry River, 4 Aug 1967, W.A. Page, 2 (JA 702) [UCLA]. Hardwar Gap, elev. 3800 ft, 1 lpd (647-20), 2 pd (647-110,111) [UCLA]. Kingston, July 1905, M. Grabham, 1 ? [USNM]. Newcastle, elev. 4000 ft, 5 Sept 1964, W.A. Page (JA 125), 1 lp? (125-10), 2 pd (125-102,106), 7 p? (125-100,101,103-105,107,114), 1 IP (125-11), 1 6, 7 2, 4 L; track to Catherines Peak, elev. 4000-4300 ft, 8 Nov 1966, O.G.W. Ber- lin, 2 pd (677-101,102), 1 p? (677-100), 5 2 (JA 679), 2 pd (680-110,111), 1 p? (680-112), 3 6, 1 P (JA 680), 1 Ipod (681-10), 1 pd (681-100), 1 6, 2 9, 2 p, 7 L(JA 681), 1 pd (682-101), 3 p? (682-100,102,103), 3 IP (682-10-12), 3 6,5 P, 13 L, 21(JA 682). Uester Road, elev. 3400-4500 ft, 8 Sept 1965, J.N. Belkin, 2 IP (337-12,13), 1 6, 1 2 (JA 337), 1 L(JA 338) [UCLA]. 9. Aedes (Howardina) grabhami Berlin, n.sp. Figs.3,7,8,21,22 TYPES: Holotype ¢ (JA 688-20) with associated pupal and larval skins and genitalia slide, near Hope Farm, Porus, Manchester, Jamaica, elev. 1000 ft, 12 Nov 1966, O.G.W. Berlin [USNM]. Al- lotype ? (JA 688-11) with associated pupal and larval skins, same data as holotype [USNM]. Para- types: 11 6 (JA 688-13,100,101,107,108,111,114-116,126,128), 14 9 (JA 688-12,15,102,103, 110,113,120-123,129-132), 1 IP (JA 688-16), 1 6,5 p, 8 L, 1 1, same data as holotype [USNM, BM, UCLA]. 1906.Howardina aureostriata Grabham, 1906:171-173. TYPES: 6, °, larva, Newcastle (St. An- drew), Jamaica, elev. 4000 ft, Sept 1905, Loscombe and Maclaverty [NE]. Aedes (Howardina) aureostriata in part of Bonne and Bonne-Wepster (1925 :370-371). Aedes aureostriata of Gowdey (1926:73). Berlin: Subgenus Howardina 43 Aedes aureostriata in part of Howard, Dyar and Knab (1917:855-859); Dyar (1918:73,80). Howardina aureostriata of Theobald (1910:609-611). Haemagogus aureostriata of Dyar and Knab (1906a:166,167). Aedes (Howardina) inaequalis in part of Thompson (1947:78; 1956:199-200); Stone, Knight and Starcke (1959:174); Belkin, Schick and Heinemann (1965:31); Forattini (1965 :399,400). Aedes inaequalis in part of Stone and Knight (1956:219). Aedes (Howardina) aurites in part of Dyar (1928:236); Edwards (1932:155); Lane (1939:99; 1953:720-722); Horsfall (1955:471). Aedes aurites in part of Hill and Hill (1948:29,42). FEMALE. Wing: 2.15 mm. Proboscis: 1.68 mm. Forefemur: 1.33 mm. Abdomen: about 2.1 mm. Essentially similar to aurites, differing mainly in the following. Head: Erect scales entirely dark. Thorax (fig.8): Mesonotal dark scales not as dense as in aurites; light lines usually distinctly narrower than in aurites, in a few specimens broader than in aurites and rarely almost completely fused; outer dorsocentral line usually distinctly shorter, not merging with lateral marginal line anteriorly, in a few specimens merging with lateral marginal line; lateral marginal line long, moderately broad, extending from side of anterior promontory to wingbase. Antealar area above paratergite predominantly with scattered golden and a few bronzy scales. Lateral scu- tellar lobes with a few narrow golden scales only. Apn with a moderate silvery patch; ssp with a distinct silvery patch. Patches on base of pra, upper and lower stp distinct, separate. Legs: Midtarsus with few white scales on segments 1,2; hindtarsus with moderate white rings on 1-3. Haltere: Stem pale, knob predominantly white but with a few dark scales dorsally. Abdomen: Sternites II, III straw yellow, [V predomi- nantly straw yellow, a few apical scales dark; rest of segments with basal yellow and distal dark scales. MALE. Wing: 2.21 mm. Proboscis: 1.75 mm. Forefemur: 1.31 mm. Essentially similar to female in color. MALE GENITALIA (fig.21). As figured; diagnostic characters as in the key. Ninth tergite lobe indistinct, with 2 strong bristles. Sidepiece length about 3.0 median width. Clasper length 0.65 of sidepiece, basal 0.33 with minute spicules; spiniform about 0.2 of clasper. PUPA (fig.21). Abdomen: 2.75 mm. Trumpet: 0.38 mm; index 4.6-4.8. Paddle: 0.69 mm. Chaetotaxy as given below, hairs moderately pigmented. Readily separated from inaequalis and stenei by the absence of lightly pigmented area on metanotum and abdominal segments uniformly lightly pigmented; from aurites by the hair 2-VI distinctly mesad or at level of 3-VI. Cephalothorax: Mesonotum and metanotum lightly pigmented. Hair 1(1), 2(2, 2-3), 3(1), 4(7, 4-8), 5(5, 4-7), 6(1, 1-2), 7(4, 3-5), 8(1, 1-3), 9(1, 1-2), 10(22, 18-32), 11(1), 12(5, 3-6). Abdomen: Lightly pigmented, progressively lighter caudad. Segment I: hair 1(12-16 primary branches, subequal in length with many secondary branches), 2(2, 1-3), 3(1), 4(8, 6-8), 5(8, 4-8), 6(1, 1-2), 7(4, 3-4), 9(1), 10€1), 11€1). Segment II: hair 0(1), 1(60, 50-74), 2(1), 3(1), 465, 4- 6), 5(1), 6(1), 7(5, 4-6), 9(1). Segment III: hair 0(1), 1(13, 8-17), 21), 3d, 1-3), 4¢4, 3-6), 5(1), 6(1), 7(4, 3-6), 8(4, 2-4), 9(1), 10(2, 2-3), 11(1), 141). Segment IV: hair O(1), 109, 8-14), 2(1), 3(4, 4-6), 4(4, 2-4), 5(1), 6(1), 7(4, 3-6), 8@2, 1-4), 9(1), 10@, 1-2), 11(1), 14(1). Segment V: hair 0(1), 1(1), 2(1), 3@, 1-3), 4(4, 4-7), 5(1), 6(1), 7(5, 4-6), 8(2, 2-3), 9(1), 10(1, 1-2), 11(1), 141). Segment VI: hair 0(1), 1(1, 1-2), 2(1, 1-2), 3(2, 1-2), 4(4, 3-4), 5(1), 6(1), 7(1), 8@, 2-3), 9€1), 10€1, 1-2), 11(1), —14(1). Segment VII: hair 0(1), 1€1), 2(1), 3(2, 1-2), 4(2), 5(1), 6(2, 2-5), 7(1), 8G, 2-4), 9(2, 2-4), 10(1), 11€1), 14(€1). Segment VIII: hair 0(1), 4(2, 1-2), 9(10, 8-11), 4d Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 14(1). Paddle: Almost without pigmentation, midrib faintly pigmented. Male genital lobe extending to 0.42 of paddle, female genital lobe to 0.35. LARVA (fig.22). Head: 0.7 mm. Siphon: 0.92 mm. Anal Saddle: 0.22 mm. -Chae- totaxy as given below, hairs lightly to moderately pigmented. Readily separated from stenei by the small and single hair 2-VII; from aurites by hair 5-C always single; from inaequalis by hair 5-C single and longer, narrow gills. Head: Width about 1.14 of length. Mental plate with a strong median tooth and 10 distinct teeth (9-12) on each side, often: a small detached tooth at base. Hair 0(1), 1(1), 3(1), 4(3, 2-5), 5(1), 6(6, 5-10), 7(3, 2-3), 8(1), 9@, 2-3), 10(2, 1-2), 11(2, 1-7), 12(@2, 2-4), 131), 14(1), 15@G, 2-4). Antenna: Length about 0.49 of head, very lightly pigmented and with a few scattered spicules. Hair 1-A at about 0.46 from base. Thorax: Stellate hairs with a few, weakly barbed branches; long hairs barbed, with attenuate apices. Prothorax: hair 0(7, 4-12), 1(6, 5-9), 2(1), 3(13, 8-19), 4(1, 1-2), 5(2, 1-2), 61), 71), 8@, 2-11), 9(2, 1-2), 10(1), 11@2, 2-4), 121), 14(13, 9-22). Mesothorax: hair 1(2, 2-13), 2(1), 3(1), 4(1), 5(1), 6(4, 2-6), 7€1), 8(5, 4-7), 9(7, 6-8), 1001), 11@, 1-2), 121), 1344, 2-4), 14(5, 4-7). Metathorax: hair 1(2, 1-12), 2(1), 3(4, 2-5), 4(2, 2-10), 5(5, 5-14), 6(1), 7(7, 6-8), 8(5, 4-8), 9(6, 6-9), 10(1), 11(2), 121), 13(4, 3-10). Abdomen: Long and stellate hairs as on thorax. Segment I: hair 1(3, 2-14), 2(6, 6-14), 3(1), 4(4, 3-4), 5(4, 4-12), 6(3), 7(1), 9(1), 10(1), 1105, 5-11), 13(4, 4-11). Segment II: hair 0(1), 1(2, 2-12), 2(5, 5-13), 31), 4(2, 2-4), 5(7, 5-14), 6(3, 2-3), 7(2, 2-3), 8(1), 9(5, 5-9), 101), 11€1), 12(1), 13(3, 2-8). Segment III: hair 0(1), 1(2, 2-3), 2(5, 5-11), 31), 4(2, 2-3), 5(7, 5-12), 6(1, 1-2), 7(6, 3-7), 8(1), 9(5, 4-9), 10€1), 11€1), 121), 13@, 2-8), 14(1). Segment IV: hair 0(1), 1(2, 1-3), 2(5, 5-12), 3(1), 4Q, 2-3), 5(6, 6-14), 6(1), 7(6, 5-7), 8(1), 9(5, 5-9), 10(1), 11€1), 121), 13(3, 3-8), 14(1). Segment V: hair O(1), 1(2, 2-3), 2(5, 5-10), 3(1), 4(4, 3-5), 5(5, 4-13), 6(1), 7(6, 3-7), 8(1), 965, 5-7), 101), 11€1), 121), 13(3, 2-8), 14(1). Segment VE: hair 0(1), 1@2, 1-3), 2(4, 4- 8), 3(1), 4(1), 5(2, 2-9), 6(1), 7(3, 2-5), 8(3, 2-3), 9(3, 2-7), 10(1), 1101), 12(1), 13, 2-11), 14(1). Segment VII: hair 0(1), 1(2, 1-2), 2(1), 31), 41), 5(3, 2-3), 6(3, 2-5), 71), 8(5, 5-10), 9(€1, 1-2), 10(1), 1101), 121), 131, 1-3), 141). Segment VIIT: Comb scales 16(15-19), moderately pigmented, elongate, fringed on sides and apex. Hair 0(1), 1(3, 3-4), 2(1), 365, 4-7), 4(1), 5(2, 2-5), 141). Siphon: Index about 6.8- 8.5; moderately pigmented, with darker basal ring. Pecten extending to about 0.44; teeth 17(14-22). Hair 1(2, 2-3), 2(1), 3-5 not visible, 6(1), 7(1), 8(2, 1-2), 9(1). Anal Segment: Saddle moderately pigmented. Gills narrow, length about 1.5 of saddle, ventral 0.8 of dorsal. Hair 1(1), 2(5, 5-8), 3(1), 4a(5, 4-7), 4b(5, 4-5), 4c(4, 3-5), 4d(4, 4-5), 4e(4, 2-6), 4f(3, 2-4). SYSTEMATICS. Although it is almost certain that grabhami represents the same taxon as aureostriatus Grabham, 1907, which is preoccupied in Aedes by aureosStria- tus (Doleschall, 1857) and aureostriatus (Leicester, 1908), I am proposing it as a new Species and not as a replacement name for aureostriatus because the types of the lat- ter are lost and therefore the identity of this nominal taxon cannot be objectively de- termined. This species was believed to be identical with inaequalis and aurites by Thompson (1956) who used the former name because of priority. A. grabhami is readily differentiated from the other species of the Aurites Group in all stages by the characters indicated in the diagnosis and keys. There is considerable variation in the ornamentation of adults of grabhami. The majority have distinct golden mesonotal lines, broader than in inaequalis and stenei and narrower than in aurites; a few have these lines narrower than in inaequalis or stenei. The most striking departure is found in a few specimens with the mesonotum almost completely golden except for a few dark scales in indistinct narrow longitu- Berlin: Subgenus Howardina 45 dinal lines; in these the hindtarsal light markings are restricted to segments 1 and 2. I have retained these specimens under grabhami on the basis of similarity in other diagnostic features but their exact taxonomic status is doubtful. They may represent a distinct species or possibly hybrids between typical grabhami and stenei which have been found in the same collections. There is also considerable variation in the chaeto- taxy of the larvae, some being much more “hairy” than others; however, the diagnos- tic characters given above apply to all the material I have seen. BIONOMICS. The immature stages of grabhami are found in both terrestrial and epiphytic bromeliads. A. stenei is frequently associated with grabhami in the same bromeliads but it is not known if the 2 species actually occupy the same axils. I did not find grabhami above 1,800 feet elevation but the original material of aureostria- tus is reported to have been collected at an altitude of about 4,000 feet. A. grabhami is, next to stenei, the most common member of the Aurites Group and has been re- ported from all parishes of Jamaica except Westmoreland, Trelawny and St. Elizabeth. Females were found to bite humans along with stenei and inaequalis and showed the same small peak of biting activity at 1700 hrs, as the latter (fig.7). DISTRIBUTION (fig.3). Endemic to the island of Jamaica where it has been re- ported from all parishes except Westmoreland, Trelawny and St. Elizabeth at eleva- tions of 100 ft to 1800 ft. Material examined: 1223 specimens; 172 6, 385 9, 373 pupae, 293 larvae; 298 individual rearings (198 pupal, 69 larval, 31 incomplete). JAMAICA. Clarendon: Belcarres, 7 Nov 1967, W.A. Page (JA 902), 1 pd (902-101), 1 p? (902- 102), 3 L [UCLA]. Hanover: Lethe, elev. 500 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson, 2 L (JA 667) [UCLA]. Manchester: Hope Farm, 8 July 1945, G.A. Thompson, 1 Ipdé (450708-2), 2 lp? (450708-9,11), 1 lp (450708-7), 1 1(450708-8); 1 Aug 1945, G.A. Thompson, 1 16 (450801-12), 1 2 (450801-12), 2 1(450801-6,8); 1 Oct 1945, G.A. Thompson, 2 Ip¢ (451001-6,9), 1 16 (451001- 1), 1 Ip (451001-14); Oct 1945, 3 L; 1 Jan 1946, G.A. Thompson, 2 lp? (460101-4,12), 16 (460101-15); 8 Jan 1946, 1 6, 1 9 [USNM]; elev. 1000-1100 ft, 10 Sept 1965, W.A. Page, 1 Ip? (354-12), 2 pd (354-105,106), 3 p? (354-103,104,107), 1 d (JA 355), 1 9 (JA 356); 12 Nov 1966, O.G.W. Berlin, 210 2 (JA 684), 2 pé (686-100,102), 1 p? (686-101), 2 Ip? (687-11,12), 5 pd (687- 100-104), 1 IP (687-10), 1 P (JA 687), 2 Ipd (688-13,20), 3 Ip? (688-11,12,15), 10 pd (688-100, 101,107,108,111,114-116,126,128), 12 p? (102,103,110,113,120-123,129,130-132), 1 IP (688- 16), 16,5 p,8L, 11(JA 688, type series) [UCLA]. Mandeville, 5 Mar 1928, Pickering, 5 9 [USNM]. Porus, elev. 900-1000 ft, 23 Oct 1966, O.G.W. Berlin and D.C. Watson, 1 lpé (630-11), 1 lp? (630- 10), 1 L (JA 630), 4 pd (631-105,106,109,112), 2 p? (631-108,113), 29,1 P,2p,1L,11(JA 631), 2 Ipd (632-10,12), 1 Ip? (632-11), 6 pd (632-101,103,104,108,112,114), 5 p? (632-100,102, 105-107), 2 IP (632-13,15), 1 P, 2 p, 19 L, 21 (JA 632); 1 Nov 1966, O.G.W. Berlin, 1 Ipé (655- 10), 6 pd (655-101,103,105-107,109), 4 p? (655-100,104,108,111), 4 IP (655-11-14),4.L,61(JA 655), 2 L (JA 656), 2 pd (658-110,114), 4 p? (658-103,107,112,113), 16,1 p, 1 L, 11(JA 658) [UCLA]. Portland: Kemney-Sherwood Forest, elev. 750 ft, 12 Aug 1967, J.N. Belkin and W.A. Page, 1 Ipé (706-11), 1 p? (706-100), 1 p (JA 706), 2 lp? (708-13,16), 1 pd (708-11), 1 Ip (708- 14), 1 @ (JA 708) [UCLA]. St. Andrew and Kingston: Hermitage Dam Road, 1 Sept 1965, W.A. Page (JA 319), 1 lp? (319-105); 17 Aug 1967, J.N. Belkin and W.A! Page, 2 lpd (723-21 ,23), 1 lp? (723-24), 1 IP (723-20), 2 p? (725-113A,114), 1 6, 1 P(JA 725); 5 July 1967, W.A. Page and IN. Belkin (JA 765), 1 pd (765-100); 25 Aug 1967, W.A. Page (JA 854), 1 Ipd (854-13), 2 po (854- 105,106), 1 p? (854-107), 2 IP (854-15,16), 2 L;5 Aug 1967, W.A. Page and J.N. Belkin (JA 892), 4 Ip (892-10,13,15,16), 3 Ip? (892-11,12,14), 2 pd (892-100,104), 3 P, 1 L [UCLA]. Kingston, M. Grabham, 1 6 [USNM]. St. Ann: Moneague, 17 Aug 1967, J.N. Belkin and W.A. Page (JA 733), 1 pd (733-105), 4 p? (733-40,90,96,114), 2 6, 1 P,6 p, 3 L, 11 [UCLA]; Faiths Pen Schoolyard, elev. 1750 ft, 31 Aug 1967, W.A. Page (JA 762), 2 pé (762-100,101), 16,19, 1P,11 [UCLA]; 2.0 mi S of Moneague near Unity, elev. 1750 ft, 25 Aug 1967, W.A. Page (JA 758), 6 pd (758-100, 103,104,109,110,113), 2 p? (758-30,101), 1 6, 2 9, 3 p [UCLA]; 29.0 mi S of Moneague, elev. 1750 ft, 30 Aug 1967, W.A. Page, 1 6, 2 9, 4 P, 31 (JA 763), 2 lpd (764-10,12), 1 Ip? (764-30), 46 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 9 pd (764-91,94,95,99 103-106,109), 5 p? (764-92,98,100,102,108), 2 d, 12,8 P, 6p, 21LQJA 764) [UCLA] ; Moneague-Claremont Road, 17 Sept 1965, W.A. Page, 1 lpd (375-12), 1 lp? (375- 13), 1 p? (375-100), 7 2 (JA 375), 1 pd (378-100), 1 p? (378-101), 1 IP (378-13), 1 Ipd (380-12), 4 Ip? (380-10,11,13,17A), 3 pd (380-101,104,105), 2 p? (380-100,107), 5 IP (380-14,15,17,18, 20) [UCLA]; Moneague-Ocho Rios Road, 10 Sept 1967, W.A. Page (JA 776), 1 p? (776-101) [UCLA]. Mt. Diablo, 17 Sept 1965, W.A. Page (JA 376), 1 Ip? (376-12); 17 Aug 1967, J.N. Belkin and W.A. Page, 6 Ipd (731-21,33,35-38), 1 lp? (731-31), 9 pd (731-90,92,93,105,108-112), 12 p¢ (731-80,81,91,94,97,99,103-107,113), 2 IP (731-30,34), 11 6,79, 2P,17 p, 24 L(VJA 731), 1 Ipd (732-10), 10 pd (732-70,94,97,100-102,104 111,112,114), 6 p? (732-90,92,93,105 ,107,108), 1 °, 1 P,3 p, 5 L, 21(JA 732), 4 lpd (733-13-16), 1 pd (733-100), 1 IP (733-11), 12 L JA 733), 1 pd (734-102), 2 p? (734-101,105), 2 P, 1 L (JA 734), 1 2 (JA 735) [UCLA] . St. Elizabeth: Darling- ton; 28 Oct 1967, W.A. Page, 1 p,4 L(JA 901) [UCLA]. St. James: Five Miles, elev. 100 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson (JA 676), 1 Ip? (676-11), 2 pd (676-115,121), 6 p? (676- 104,107,112,113,116,123), 1 IP (676-17), 1 3, 1 2, 3 p, 2 L [UCLA]. Montego Bay, elev. 500 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson (JA 669), 1 IP (669-11) [UCLA]. Reading, elev. 200 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson, 1 1 (JA 670) [UCLA]. St. Mary: Broadgate, elev. 500 ft, 27 Oct 1966, O.G.W. Berlin and D.C. Watson (JA 643), 2 lpd (643-33,43), 1 lp? (643-37), 1 L,11 [UCLA]. Leith Hall, 13 Nov 1966, O.G.W. Berlin and D.C. Watson, 4 L (JA 689) [UCLA]. Windsor Forest, elev. 750 ft, 30 Oct 1966, O.G.W. Berlin and D.C. Watson, 1 Ip? (650-22), 4 pd (650-100-102,105), 2 IP (650-20,23), 1 6, 6 P, 3L (JA 650), 1 IP (651-34), 2 Ip? (652-10,12), 4 pd (652-100,102,103,110), 4 p? (652-101,104,108,111), 2 IP (652-14,15), 11 L, 11(JA 652) [UCLA]. Parish not specified: 1944, R.B. Hill, 1 6; 1 Jan 1946, G.A. Thompson, 7 L; M. Grabham, 1 Ip; G.A. Thompson, 1 6, 2 9? [USNM]; W.A. Page, 2 pd (771-101,106), 4 L (JA 771), 2 Ip? (772-33, 34), 8 pd (772-90,93,95,97,99,106,110,114), 11 p? (772-91,92,96,98,101 ,103,104,107,108,111, 113), 1 lp (772-30), 3 6,6 9,5 P, 11 p, 40 L, 11(JA 772) [UCLA]. 10. Aedes (Howardina) inaequalis (Grabham) Figs.3,7,8,23,24 1907.Howardina inaequalis Grabham, 1907:25-26. TYPE: Lectotype 6 with genitalia slide, Kings- ton (St. Andrew), Jamaica, date not specified, M. Grabham [USNM; selection of Stone and Knight 1956:219]. Aedes (Howardina) inaequalis in part of Thompson (1947:78; 1956:199-200); Stone, Knight and Starcke (1959:174); Belkin, Schick and Heinemann (1965:31); Forattini (1965:399,400); Por- ter (1967:38). Aedes inaequalis in part of Stone and Knight (1956:219). Howardina inaequalis of Theobald (1910:611); Surcouf and Gonzalez-Rincones (1911:141). Aedes (Howardina) aurites in part of Dyar (1928:236); Edwards (1932:155); Lane (1939:99; 1953:720-722); Horsfall (1955:471). Aedes aurites in part of Hill and Hill (1948:29,42). Aedes (Howardina) aureostriata in part of Bonne and Bonne-Wepster (1925:370-371). 7 Aedes aureostriata in part of Howard, Dyar and Knab (1917:855-859); Dyar (1918:73,80). FEMALE. Wing: 2.37 mm. Proboscis: 1.93 mm. Forefemur: 1.49 mm. Abdomen: about 2.1 mm. Similar to aurites, differing primarily in the following. Head: Erect scales dark. Thorax (fig.8): Mesonotal dark scales not dense as in aurites; mesonotal light lines narrow, faintly golden, distinctly narrower than dark lines; outer dorsocen- tral line shorter, not merging with lateral marginal line anteriorly. Lateral marginal line narrow from side of anterior promontory to wingbase. Antealar area above para- tergite with scattered dark scales only. Lateral scutellar lobes with a few narrow gold- en scales. Apn with a restricted silvery patch, distinctly smaller than in aurites and Berlin: Subgenus Howardina 47 grabhami; ssp with a distinct silvery patch; patches on base of pra, upper and lower stp distinct, separate. Legs: Midtarsus with a few white scales on segments 1,2; hind- tarsus with broader white rings on 1-3. Haltere: Stem pale, knob predominantly white, with a few dark scales dorsally. Abdomen: Basomedian tergal straw yellow patches on V-VII. | MALE. Wing: 2.36 mm. Proboscis: 2.01 mm. Forefemur: 1.58 mm. Essentially similar to female in coloration. MALE GENITALIA (fig.23). Diagnostic characters as in the key. Ninth tergite lobe not prominent, with 2,3 bristles. Sidepiece length about 2.5 median width. Clasper length about 0.55 of sidepiece: spiniform short, about 0.25 of clasper. PUPA (fig.23). Abdomen: 3.08 mm. Trumpet: 0.46 mm; index 5.0-5.6. Paddic: 0.68 mm. Chaetotaxy as given below, hairs lightly to moderately pigmented. Very similar to stenei but apparently distinguishable by the moderately pigmented abdom- inal tergite II and segment III only slightly darkened in the middle, without distinct dark stripe. Readily separated from aurites and grabhami by the lightly pigmented area on the metanotum. Cephalothorax: Mesonotum moderately pigmented between trumpets; lateral area slightly darker; metanotum with a small to moderate, lightly pigmented area in the middle of lateral portion. Hair 1(1), 2(2, 1-3), 3(1, 1-3), 4(4, 2-7), 5(6, 3-7), 6(1, 1-2), 7(4, 3-6), 8(1, 1-2), 9(2, 2-3), 10(15, 6-15), 111), 12(4, 2- 5). Abdomen: Tergite I moderately pigmented except for usual membranous areas; tergites II,III moderately pigmented; other segments lighter. Segment I: hair 1(11-14 primary branches, subequal in length, each with many secondary branches), 2(1, 1-2), 3(1), 405, 4-7), 5(5, 4-6), 6(1), 7(3, 2-3), 9(1), 10(1, present in some), 11(1, present in some). Segment II: hair 0(1), 1(40, 36-56), 2(1), 3(1), 4(4, 4-8), 5(1), 6(1), 7(4, 3-4), 9(1), 10(2, 2-3, present in a few). Segment III: hair 0(1), 109, 9-20), 2(1), 31), 4(4, 2-4), 5(1), 6(1), 7(4, 2-5), 8(3, 2-3), 9(1), 10(2), 11(1), 141). Segment IV: hair O(1), 1(6, 3-14), 2(1), 3(4, 3-7), 4(2, 2-3), 5C1, 1-2), 6(1), 7(3, 2-4), 8@2, 1-2), 9C1), 10(2, 1-2), 11€1), 14(1). Segment V: hair 0(1), 1(1, 1-4), 2(1), 3@, 1-3), 4(6, 3-7), S(1, 1-2), 6(1), 765, 3-7), 8@, 1-3), 9(1), 10(1, 1-2), 11€1), 141). Segment VI: hair O(1), 1€1, 1-3), 2(1), 31, 1-3), 4(3, 2-7), 5C1, 1-2), 6(1, 1-2), 7(1, 1-2), 8@3, 2-4), 9(1), 10(€1), 111), 14(1). Segment VII: hair 0(1), 1(1), 21), 31), 4@, 1-3), 5(1), 6(3, 2-4), 7(1, 1-3), 8(3, 1-5), 9(4, 3-5), 10(€1), 11(1), 14(1). Segment VIII: hair 0(1), 4(2, 1-2), 9, 8-12), 14(1, 1-2). Paddle: Uniformly lightly pigmented, midrib slight- ly darker; external margin smooth; hair 1-P slender. Male genital lobe extending to 0.56 of paddle, female genital lobe to 0.36. LARVA (fig.24). Head: 0.7 mm. Siphon: 0.9 mm. Anal Saddle: 0.22 mm. Readily separated from other species by the short, broad unequal gills and hair 5-C usually double. Head: Width about 1.12 of length. Mental plate with a strong median tooth and 11 distinct teeth (11-12) on either side, often a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 4(3, 2-4), 5@2, 1-2), 6(9, 8-11), 7(4, 3-4), 8(1), 9(2, 2-3), 10(1), — 1102, 2-5), 12(1, 1-2), 131), 14(1), 153, 2-3). Antenna: Length about 0.46 of head; pigmentation very light; shaft with very few minute scattered spicules. Hair 1-A about 0.5 from base. Thorax: Stellate hairs with a few unequal branches; long hairs barbed and with attenuate apices. Prothorax: hair 0(4, 2-8), 1(3, 3-7), 2(1), 3(5, 4 6), 4€1, 1-2), 5(2), 6(1), 701, 1-2), 8(3, 2-4), 9(1, 1-2), 10(1), 11(2, 2-4), 12Q), 14(10, 9-12). Mesothorax: hair 1(2, 2-5), 2(1), 3(1), 4(1), 5(1), 6(4, 4-5),-7(1), 8(6, 5-8), 9(7, 5-8), 10(1), 11@2, 2-3), 12(1), 13(11, 7-11), 14(5, 4-5). Metathorax: hair 1(2, 2-4), 2(1), 3(4, 3-5), 4(2, 2-4), 5(5, 4-6), 6(1), 7(6, 5-7), 8G, 5-9), 9G, 5-7), 10(1), 11(2, 1-2), 12(1), 13(4, 1-5). Abdomen: Long and stellate hairs as on thorax. Segment I: hair 1(3, 2-7), 2(5), 3(1), 4(3, 3-4), 5(4, 3-6), 6(3, 2-3), 7(1, 1-2), 9(1), 48 : Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 10(1), 11(6, 5-8), 13(3, 2-7). Segment II: hair 0(1), 1(2, 2-6), 2(5, 4-6), 3(1), 4@, 2- A), 5(6, 4-8), 6(3, 2-3), 7(3, 2-4), 81, 1-2), 9(5, 4-6), 101), 11(1), 121), 13@, 2-4). Segment III: hair 0(1), 1(2, 2-4), 2(5, 5-6), 3(1), 4(2), 5(5, 5-8), 6(1), 765, 3-6), 8(1), 9(5, 4-5), 10(1), 11(1), 12(1), 13(4, 2-8), 14(1). Segment IV: hair 0(1), 1@, 2-4), 2(6, 5-6), 3(1), 4(2), 5(7, 5-9), 6(1), 7(5, 3-6), 8(1), 9(5, 4-5), 10(1), 11€1), 121), 13(5, 3-10), 14(1). Segment V: hair 0(1), 1(3, 2-4), 2(5, 5-6), 3(1), 4(@2, 1-3), 5(6, 5- 9), 6(1), 7(5, 3-5), 8C1, 1-2), 965, 4-6), 10(1), 111), 12(1), 13(4, 2-9), 14C1). Seg- ment VI: hair 0(1), 1(2, 2-3), 2(4, 4-5), 3(1), 4(1), 5(4, 2-6), 6(1), 7(3, 2-4), 8(3, I- 3), 9(3, 2-4), 10(1), 11(1), 121), 13(3, 24), 14(1). Segment VII: hair 0(1), 1(2), I(1), 31), 401, 5C, 2-3), 64, 2-5), 70). 8C7; 6-9); 9G,.24), 1001), 1 LG), IP2c1), 13(2), 14(1). Segment VIII: Comb scales 16(14-18) in number; individual scale mod- erately pigmented, elongate, fringed on sides and apex. Hair 0(1), 1(4, 3-5), 2(), 3(5, 5-6), 4(1), 5(3, 3-5), 14(1). Siphon: Index about 4.25 to 5.3; moderately pig- mented but with darker basal ring. Pecten extending to about 0.4, 19(16-25) teeth; individual tooth narrow, long, with minute spinules in distal half. Hair 1(2), 2(1), 3-5 not visible, 6(1), 7(1), 8(1), 9(1). Anal Segment: Saddle moderately pigmented. Gills as long as or slightly shorter than saddle length, broad, unequal, ventral 0.64 of dor- sal. Hair 1(1), 2(5, 4-5), 3(1), 4a(4, 3-4), 4b(4), 4c(4, 3-4), 4d(4), 4e(3), 4£(3, 1-3). SYSTEMATICS. Contrary to the opinion of Thompson (1956:199), inaequalis is a species distinct from aureostriatus (now grabhami) and aurites. The adults of these 3 forms are readily distinguished from one another by the characters mentioned in the diagnoses and keys but I have not been able to find any reliable characters to sep- arate inaequalis from Thompson’s stenei which, however, I consider to be also a dis- tinct species. The larva of inaequalis is clearly differentiated from the other members of the group but the pupa is very similar to that of stenei. Furthermore, an ecological barrier separates inaequalis from the rest of the group as this species is never found breeding in bromeliads to which the other species are confined. A. inaequalis is found primarily at low elevations and seldom above 1,500 feet. I have seen specimens of in- aequalis only from the parishes of Clarendon, Kingston, Manchester, Portland, St. Andrew, St. Catherine, St. Mary and St. Thomas. This species may be more widely distributed and more common than the records indicate because these are based on immature stages only and these breeding sites have not been as widely sampled as those of the other species of the group. : BIONOMICS. A. inaequalis has been found breeding primarily in treeholes and cut and broken bamboo but has also been taken occasionally in rockholes and crab- holes. There are no records of this species breeding in bromeliads. Since adults cannot be distinguished at present from stenei, it is not known if females of inaequalis bite humans but it is possible that some females of this species are included in the stenei- inaequalis material obtained in the 24-hour biting-landing survey (fig.7). DISTRIBUTION (fig.3). Endemic to the island of Jamaica where it has been re- ported from the parishes of Clarendon, Kingston, Manchester, Portland, St. Andrew, St. Catherine, St. Mary and St. Thomas from near sea level to about 1000 ft. Mate- rial examined: 574 specimens; 55 6, 81 2, 99 pupae, 339 larvae; 83 individual rear- ings (47 pupal, 28 larval, 8 incomplete). JAMAICA. Clarendon: May Pen, 10 Sept 1965, rockholes, J.N. Belkin (JA 352), 6 pd (352- 100A,102,104,106,107,109), 6 p? (352-32,90,91 93,108,111), 1 IP (352-12), 6 6, 8 2, 1 P, 8 p, 37 L; 6 Sept 1966, rockholes, D.C. Watson (JA 602), 1 pd (602-101), 1 p? (602-100), 5 6, 7 9; 1 Sept 1966, rockholes, D.C. Watson, 2 6 (JA 603) [UCLA] . Manchester: Hope Farm, elev. 900- 1000 ft, 23 Oct 1966, treehole, O.G.W. Berlin and D.C. Watson (JA 634), 2 Ipd (634-10,17), 5 Ip? — (634-11-13,15,19), 2 pd (634-102,104), 3 p? (634-100-102), 4 IP (634-14,16,18,20), 1 L, 11 Berlin: Subgenus Howardina 49 [UCLA]. Portland: Kemney-Sherwood Forest, elev. 750 ft, 13 Aug 1967, bamboo, J.N. Belkin and W.A. Page, 1 lpd (713-10), 1 pd (713-100), 16 L(JA 713), 2 Ipd (715-10,11), 2 pd (715-100, 102) [UCLA]. St. Andrew and Kingston: Constant Springs, 16 Nov 1966, treehole, O.G.W. Berlin and D.C. Watson, 6 L (JA 696) [UCLA]. Ferry Quarry, 13 Sept 1965, crabhole, W.A. Page, 1 ? (JA 369) [UCLA]. Hermitage Dam Road, 20 Aug 1964, rockhole, L. Paton (JA 109), 2 Ipd (109- 10,11), 1 pd (109-101), 1 p? (109-100), 4 3, 1°,1P,2L;22 Aug 1965, treehole, W.A. Page (JA 280), 1 pd (280-102), 1 6; 10 Aug 1965, treehole, W.A. Page (JA 288), 2 Ip? (288-10,15), 1 pd (288-102), 1 6; 13 Aug 1965, bamboo, W.A. Page, 1 Ipd (290-14), 3 Ip? (290-10,11,13),2L(JA 292), 25 L (JA 293), 6 L(JA 294), 1 Ip? (295-23), 26, 19,10 L(JA 296), 1 L(JA 298),12(JA 299), 1 p? (300-20), 2 L (JA 300), 17 L (JA 302), 1 lp? (304-22), 3 L (JA 307), 24 L(JA 308), 1d, 19, 1 P (JA 309); 7 Sept 1965, J.N. Belkin, 1 9, 2 L(JA 330), 3 L(JA 331); 29 Oct 1965, treehole, W.A. Page, 1 pd (390-100A), 2 p? (390-101,102), 1 p? (397-105); 13 July 1967, tree- hole, W.A. Page (JA 826), 1 p? (826-20) [UCLA]. Mt. Salus, 17 Sept 1967, bamboo, W.A. Page, 123 L (JA 780) [UCLA]. Temple Hall, elev. 500 ft, 20 Aug 1965, bamboo, W.A. Page, 1 p? (250- 102), 1 6,1 9,1 P,5 L(JA 250), 1 Ipd (252-35), 4 Ip? (252-102,105,107,108), 16,1 p,6L,11 (JA 252); 25 Aug 1965, bamboo, D.C. Watson (JA 251), 1 pé (251-107), 1 IP (251-60), 1,11 L [UCLA] . Waterworks, 7 Sept 1965, treehole, J.N. Belkin (JA 327), 1 p? (327-104), 19,1 p,1L; 3 Dec 1965, treehole, W.A. Page (JA 408), 1 IP (408-14) [UCLA]. St. Catherine: Harkers Hall, elev. 750 ft, 12 July 1967, bamboo, W.A. Page (JA 822), 2 lp? (822-12,14), 1 IP (822-10), 1 , 1 p, 5 L [UCLA]. Rio Cobre Dam, elev. 100 ft, 10 Sept 1965, treehole, J.N. Belkin (JA 351), 1 lp? (351-30), 1 p? (351-105), 1 IP (351-31), 1 L; 17 Sept 1965, ditch, D.C. Watson, 1 6 (JA 372), tree- hole, 1 6,3 2, 1 p(JA 373) [UCLA]. Spanish Town, 9 Sept 1965, treehole, D.C. Watson (JA 605), 1 pd (605-102) [UCLA]. St. Mary: Prospect, 3 Nov 1964, rockhole, W.A. Page, 1 6,19,1p(JA 173) [UCLA]. St. Thomas: Folly, elev. 50 ft, rockhole (JA 103), 4 p? (103-102,103,105,108), 1? [UCLA] . Parish not specified: Jan 1940, bamboo, 1 6, 2 ? [USNM]. 11. Aedes (Howardina) stenei Thompson Figs 3,7 .6.22,20 1956.Aedes stenei Thompson, 1956:196-199. TYPE: Holotype 6 (45090211) with associated lar- val skin and male genitalia slide, near Hope Farm, just above Porus, on road to Mandeville, Manchester (as Clarendon), Jamaica, elev. about 1000 ft, Sept 1945, G.A. Thompson [USNM]. Aedes (Howardina) stenei of Stone, Knight and Starcke (1959:174); Belkin, Schick and Heine- mann (1965:31); Forattini (1965 :471); Porter (1967:38). FEMALE. Wing: 2.57 mm. Proboscis: 1.66 mm. Forefemur: 1.58 mm. Abdomen: about 2.02 mm. Essentially similar to inaequalis from which it cannot be separated (fig.8). | _ MALE. Wing: 2.38 mm. Proboscis: 2.1 mm. Forefemur: 1.58 mm. Essentially sim- ilar to female in coloration. MALE GENITALIA (fig.25). Diagnostic characters as in the key. Ninth tergite lobe indistinct, with 2,3 strong bristles. Sidepiece length about 2.5 median width. Clasper about 0.6 of sidepiece length, basal 0.33 with small spicules; spiniform about 0.25 of clasper. PUPA (fig.25). Abdomen: 2.24 mm. Trumpet: 0.38 mm; index about 5.0. Paddle: 0.68 mm. Chaetotaxy as given below, hairs lightly to moderately pigmented. Very similar to inaequalis but distinguished by the deeply pigmented abdominal tergite II and by the presence of a distinct narrow median longitudinal dark stripe on tergite III. Readily separated from aurites and grabhami by the 2-branched hair 9-C and the lightly pigmented area on the metanotum. Cephalothorax: Mesonotum moderately 50 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 pigmented between trumpets, lateral areas slightly darker; metanotum with a lightly pigmented area in the middle of lateral portion. Hair 1(1), 2(2, 2-3), 3(1), 405, 4-7), 5(5, 4-9), 6(1, 1-2), 7(4, 3-9), 8(1, 1-2), 9@2, 2-3), 10(10, 7-18), 11(1), 12(5, 3-7). Abdomen: Tergite I moderately pigmented except for usual membranous areas; ter- gite II strongly pigmented; tergite III with distinct narrow, median, dark longitudinal stripe; pigmentation progressively lighter caudad. Segment I: hair 1(12-17 primary branches, subequal in length each with several secondary branches), 2(1, 1-2), 3(1), 4(6, 4-8), 5(4, 3-8), 6(1), 7(3, 1-3). 9(1), 10(1), 11(1). Segment IT: hair 0(1), 1(46, 36-72), 2(1), 3(1), 4(5, 3-6), 5(1), 6(1), 7(4, 2-5), 9(1), 10€1, 1-4, present in some). Segment III: hair 0(1), 1(16, 7-23), 2(1), 3(1, 1-2), 4(3, 2-4), 5(1, 1-2), 6(1), 7(3, 2- 4), 8(2, 2-4), 9(1), 10(2, 1-2), 111), 14(1). Segment IV: hair 0(1), 1(10, 4-19), 21), 3(5, 3-6), 4(3, 1-4), 5(1, 1-3), 6(1), 7(@2, 2-4), 8(2, 1-3), 9(1), 10@, 1-2), 11(1), 14(1). Segment V: hair 0(1), 1(2, 1-7), 2(1), 3@, 1-3), 4(4, 3-6), 51, 1-2), 6(1), 7(4, 2-5), 8(2, 1-3), 9(1), 10(1, 1-2), 11€1), 14(1). Segment VI: hair 0(1), 1(2, 1-4), 21), 3@, 1-2), 4(3, 2-5), 5(1, 1-4), 6(1, 1-2), 7(1), 8(2, 2-4), 9(1), 10(€1), 11€1), 14(1). Seg- ment VII: hair 0(1), 1(1), 21), 31, 1-2), 4(2, 1-2), 5(1), 6(3, 2-4), 7(1), 8(4, 3-5), 9(5, 3-6), 10(1), 11€1), 14(1). Segment VIII: hair 0(1), 42, 1-2), 9(10, 8-13), 14(1). Paddle: Uniformly very lightly pigmented, midrib slightly darker; external margin smooth. Male genital lobe extending to 0.42 of paddle, female genital lobe to 0.38. LARVA (fig.26). Head: 0.7 mm. Siphon: 0.95 mm. Anal Saddle: 0.27 mm. Read- ily separated from other members of the group by the strong, 4-7-branched hair 2- VII. Head: Width about 1.02 of length. Mental plate with a strong median tooth and 11 distinct teeth (9-11) on each side, often a detached tooth at base. Hair 0(1), 1(1), 3(1), 4(5, 3-5), 5(4, 3-5), 614, 11-14), 7(5, 4-7), 8(1), 9(2, 2-3), 10(1), 11(6, 3-8), 12(2, 1-2), 13(1), 141), 15(2, 2-3). Antenna: Length about 0.45 of head; pigmenta- tion very light; shaft with a few scattered spicules. Hair 1-A about 0.5 from base. Thorax: Stellate hairs with a few, unequal, weakly barbed branches; long hairs barbed and with attenuate apices. Prothorax: hair 0(12, 10-14), 1(9, 7-11), 2(1), 3(14, 8- 16), 4€1, 1-2), 5(2, 2-3), 6(1), 7(2, 2-3), 8(6, 6-10), 9(1, 1-2), 10(1), 11(3, 3-5), 12(1), 14(10, 8-14). Mesothorax: hair 1(5, 3-9), 2(1), 3(1), 4(1), 5(1), 6(5, 4-5), 7(1), 8(6, 5-7), 9(8, 7-10), 10(1), 11(2, 1-2), 12(1), 13(14, 10-14), 14(4, 4-9). Meta- - thorax: hair 1(2, 2-8), 2(1), 3(3, 3-5), 4(8, 5-9), 5(8, 6-11), 6(1), 7(7, 7-9), 8(6, 4-9), 9(8, 6-8), 10(1), 11@2, 1-3), 12(1), 13(11, 5-13). Abdomen: Stellate hairs as on tho- rax. Segment I: hair 1(5, 5-12), 2(8, 5-9), 3(1), 4(3, 2-4), 5(8, 7-11), 6(3), 7(1), 9(1), 10(1), 11(8, 7-9), 13(10, 3-12). Segment II: hair 0(1), 1(3, 2-9), 2(7, 3-8), 3(1), 4(3, 2-3), 5(9, 7-11), 6(3), 7(2, 2-3), 8(1, 1-2), 9(6, 5-7), 10(1), 11(1), 12(1), 13(8, 5-12). Segment III: hair 0(1), 1(3, 2-5), 2(7, 5-8), 3(1), 4(2), 5(10, 9-11), 6(1, 1-2), 7(6, 4- 7), 8(1), 9(6, 5-7), 101), 1101), 121), 1309, 5-13), 14(1). Segment IV: hair 0(1), 103, 2-4), 2(7, 6-8), 3(1), 4(2), 5(9, 9-12), 6(1, 1-2), 7(5, 4-6), 8(1), 9(6, 5-6), 10(1), 11(1), 12(1), 13(10, 7-11), 14(1). Segment V: hair 0(1), 1(2, 2-3), 2(6, 5-7), 3(1), 4(3, 2-4), 510, 9-11), 6C1, 1-2), 7(5, 4-6), 8(1), 9(6, 5-6), 10(1), 111), 12(1), 13(8, 4-11), 14(1). Segment VI: hair 0(1), 1(2, 2-3), 2(6, 5-7), 3(1), 4(1, 1-2), 5(7, 5-10), 6(1), 7(2, 2-3), 8(3, 2-3), 9(5, 4-6), 10(1), 11(1), 12(1), 1309, 6-11), 14(1). Segment VII: hair O(1), 1(2, 2-4), 2(5, 4-7), 3(1), 4(1), 5(2, 2-3), 6(2, 2-3), 7(1), 8(5, 3-7), 9(4, 4-6), 10(1), 11(1), 121), 13(6, 3-6), 14(1). Segment VIII: Comb scales 18(16- 19) in number, individual scale moderately pigmented, elongate and fringed on sides and apex. Hair 0(1), 1(4, 4-6), 2(1), 3(6, 5-7), 4(1), 5(4, 3-5), 14(1). Siphon: Index about 5.8-6.7; moderately pigmented, with darker basal ring. Pecten extending to about 0.43; 19(19-24) narrow, long teeth. Hair 1(2, 2-3), 2(1), 3-5 not visible, 6(1), 7(1), 8(2, 1-2), 9(1). Anal Segment: Saddle moderately pigmented. Gills subequal; Berlin: Subgenus Howardina 51 moderate, less than 2.0 of saddle length. Hair 1(1, 1-2), 2(7, 5-8), 3(1), 4a(4, 4-5), 4b(4, 3-4), 4c(4, 3-4), 4d(4), 4e(4, 3-4), 4£(3, 3-4). SYSTEMATICS. A. stenei appears to be most closely related to inaequalis. | have not found any reliable characters to separate the adults of these 2 species and their pupae are also very similar but apparently differentiated by the features mentioned in the diagnoses and key. The larva of stenei is readily separated from inaequalis and the other members of the group. There appears to be a strong ecological barrier be- tween stenei, which breeds exclusively in bromeliads, and inaequalis, which is pri- marily a treehole breeder and has never been found in bromeliads. A. stenei has been collected most frequently at elevations of about 1,000 feet and less commonly down to 100 feet and as high as 2,500 feet. Its most common associate is grabhami; as in- dicated under the latter there is a suggestion of occasional hybridization between these 2 forms. A. stenei is the most common and widespread species of the group and has been reported from all the parishes of Jamaica except Clarendon, Trelawny and Westmoreland. BIONOMICS. A. stenei has been found breeding only in terrestrial and epiphytic bromeliads. Although the females of stenei and inaequalis cannot be separated with certainty, I believe that the majority of the stenei-inaequalis specimens collected in the 24-hour biting-landing study reported in fig.7 were probably stenei. These showed a small peak of activity at 1700 hrs. DISTRIBUTION (fig.3). Endemic to the island of Jamaica where it has been re- ported from all the parishes except Clarendon, Trelawny and Westmoreland at eleva- tions of 100 to 2500 ft. Material examined: 1486 specimens; 226 6, 380 2, 492 pu- pae, 388 larvae; 371 individual rearings (227 pupal, 102 larval, 42 incomplete). JAMAICA. Hanover: Lethe, elev. 500 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson (JA 666), 2 p? (666-100,101), 2 p, 2 L [UCLA]. Manchester: Hope Farm, 2 Sept 1945, G.A. Thompson, 1 Ip? (450902-2); 2 Oct 1945, G.A. Thompson, 2 Ip? (451002-10,13), 1 lp? paratype (451002-3), 1 Ipd, paratype (451002-2); 1 Jan 1946, G.A. Thompson, 1 Ipd, paratype (460101-20), 1 lp? (460101-18) [USNM]; elev. 900-1000 ft, 18 Aug 1965, W.A. Page, 1 lpd (240-62), 2 Ip? (240-63, 64), 1 IP (240-65), 1 Ip? (241-62), 7 pd (241-101,103,106-108,110,111), 1 p? (241-114), 1 IP (241-60), 2 pd (242-62,100), 4 p? (242-61,103,105,106); 10 Sept 1965, W.A. Page, 1 lp? (355- 10), 1 p? (355-100), 2 6, 2 2 (JA 355), 2 2 (JA 356); 2 Nov 1966, O.G.W. Berlin, 3 Ipd (661-10, 11,16), 2 pd (661-100,104), 3 p? (661-101-103), 4 IP (661-12-15), 19,2 p,61(JA 661), 3 2(JA 663A); 12 Nov 1966, 0.G.W. Berlin, 90 ? (JA 684), 3 P, 5 L(JA 685), 3 lpdé (688-14,17,19), 2 lp? (688-10,18), 9 pd (688-105,106,112,117-119,124,125,127), 1 p? (688-109), 5°, 1 p,91(JA 688) [UCLA]. Porus, elev. 900-1000 ft, 23 Oct 1966, O.G.W. Berlin and D.C. Watson, 1 lp? (630-12), 1 IP (630-13), 1 6, 1 p, 3 L(JA 630), 2 Ipd (631-13,14), 4 lp? (631-10,11,16,17), 4 pd (631-100, 101,104,107), 6 p?(631-102,103,110,111,114,115), 5 IP (631-18-22), 2d, 1 9, 1 p(JA 631), 1 Ipd (632-14), 3 pd (632-109-111), 1 p? (632-113), 1 IP (632-16), 1 6, 19,2P,2 p, 5 L, 31; 1 Nov 1966, O.G.W. Berlin, 1 p? (655-102), 1 1(JA 655), 3 lp? (658-10,11,13), 6 pd (658-100-102,106, 109,111), 3 p? (658-104,105,108), 1 IP (658-12), 1 9, 2 p, 21 [UCLA]. Portland: Kemney-Sher- wood Forest, elev. 750 ft, 13 Aug 1967, J.N. Belkin and W.A. Page, 2 p? (706-12,103), 146,19 (JA 706), 1 Ip? (708-10), 1 6, 1 9, 2 p, 21(JA 708), 2 L(JA 710), 2 p(JA 711) [UCLA]. St. An- drew and Kingston: Hermitage Dam Road, 1 Sept 1965, W.A. Page (JA 319), 1 pd (319-102); 29 Oct 1965, W.A. Page (JA 392), 3 pd (392-103,107,112), 2 p? (392-108,110); 15 Aug 1967, JN. Belkin and W.A. Page, 1 lp? (723-25), 2 p? (723-100,101), 1 IP (723-22), 1 9, 2P (JA 723), 2 Ipd (725-22,23), 2 Ip? (725-11,12), 7 pd (725-94,101,102A,103A,108,111,112A), 9 p? (725-100,103, 105B,106A,107A,110,111A,113,114), 2 IP (725-20,21), 3 ?, 1 P, 5 p (JA 725); 25 July 1967, W.A. Page, 3 pd (854-100,101,103), 1 p? (854-104), 1 Ip (854-12), 1 P, 11(JA 854), 1 P(JA 855) [UCLA]. Kingston, 15 Sept 1906, M. Grabham, 1 6 [USNM]. Mavis Bank, M. Grabham, 1 6 [USNM]. Mt. Salus Road, 11 Sept 1967, W.A. Page (JA 779), 2 pd (779-102,106), 1 p? (779-103) [UCLA] . Newcastle, elev. 2500 ft, 29 Oct 1966, O.G.W. Berlin, 3 L (JA 648) [UCLA]. Shooters oo Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Hill, elev. 1800 ft, 3 Dec 1946, G.A. Thompson, 2 ¢ [IJ]. Stony Hill, 16 Sept 1946, G.A. Thomp- son, 1 ? [IJ]. Temple Hall, elev.500 ft, 27 Oct 1966, O.G.W. Berlin and D.C. Watson, 3 L (JA 646) [UCLA]. St. Ann: Moneague, 1943, R.B. Hill, 1 9 [USNM]; 9 Sept 1966, D.C. Watson, 1 d,1¢ (JA 607); 17 Aug 1967, J.N. Belkin and W.A. Page (JA 733), 2 lpd (733-10,15), 9 Ip? (733-11-14, 16-19,30), 7 pd (733-90,94,97,104,107,110,113), 12 p? (733-31 ,41,92,93 ,99-102,106,111,112), 12 6,19 9,2 P, 18 p, 11 L, 31; 6 Aug 1967, W.A. Page (JA 771), 1 p? (771-102), 1 L [UCLA] ; Faiths Pen Schoolyard, elev. 1750 ft, 30 Aug 1967, W.A. Page (JA 762), 1 Ipd (762-11), 2 Ip? (762-12,13), 1 19 (762-14), 1 pd (762-103), 1 p? (762-102), 1 P, 9 L, 1 1 [UCLA]; 2.0 mi S of Moneague near Unity, elev. 1750 ft, 26 Aug 1967, W.A. Page (JA 758), 1 lp? (758-23), 4 p? (758- 31,32,108,111), 5 p? (758-105-107,112,114), 8 6, 6 2,5 p, 15 L [UCLA] ; 29.0 mi post on Mon- eague Road, elev. 1750 ft, 30 Aug 1967, W.A. Page, 1 Ipd (763-20), 6 Ip? (763-12,14,21-23 30), 8 pd (763-90,94,96-98 100,107,111), 6 p? (763-95 ,99,101-103,105), 2 6,12 9,11 P, 19 p, 18 L, 1 1(JA 763), 1 lpd (764-31), 1 Ip? (764-34), 5 pd (764-32,96,112-114), 4 p? (764-90,93,107,111), 1 lp (764-33),6 6,8 9,5 P,5 p,1L,11 [UCLA]; Moneague-Claremont Road, 17 Sept 1965, W.A. Page, 1 pd (380-106), 1 p (JA 380), 1 Ip? (381-12), 1 pd (881-100) [UCLA]; Moneague-Ocho Rios Road, 10 Sept 1967, W.A. Page (JA 776), 1 pd (776-100) [UCLA]. Mt. Diablo, 17 Sept 1965, W.A. Page, 2 Ipd (375-11,14), 2 p? (375-10,100), 2 lp (375-1522), 8 9, 21(JA 375), 3 Ipd (376- 10,11,13), 2 lp? (376-15,16), 2 pd (376-100,102), 3 P (JA 376); 17 Aug 1967, J.N. Belkin and W.A. Page, 1 Ipd (731-22), 2 Ip? (731-20,23), 1 pd (731-96), 1 p? (731-101), 16,2?,2p,3 LGA 731), 2 lp? (732-30,31), 4 pd (732-95,103,109,110), 2 p? (732-32,98), 1 IP (732-33), 2 P, 2 p, 7 L, 11(JA 732), 2 lp? (734-10,11), 4 pd (734-103 ,104,106,109), 3 p? (734-100,107,108), 1 IP (734- 12),2 9,1 p, 22 L(JA 734); 7 Sept 1967, W.A. Page, 1 lp? (779-10), 1 p? (774-107), 4 L (JA 774) _ [UCLA]. St. Catherine: Bog Walk, elev. 500 ft, 12 July 1967, W.A. Page (JA 820), 1 pd (820-11) [UCLA]. St. Elizabeth: Maggotty, elev. 500 ft, 15 Sept 1967, W.A. Page (JA 791), 1 L [UCLA]. St. James: Five Miles, elev. 100 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson (JA 676), 1 lp? (676-12), 4 pd (676-102,103,105,124), 8 p? (676-100,101,106,108,109,117,122,125), 4 d, 2 @, 2 L [UCLA]. Montego Bay, elev. 500 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson (JA 669), 1 lpd (669-10), 1 lp? (669-11), 2 pd (669-100,102), 1 p? (669-101), 3 L, 11 [UCLA]; 16 Jan 1940, 26,2 2;17 Jan 1940, 2 6, 3 9? [USNM]. Reading, elev. 200 ft, 6 Nov 1966, O.G.W. Berlin and D.C. Watson, 3 pé (670-100-102), 1 p? (670-103), 3 P, 3 p, 5 1(JA 670), 2 lp? (671-10,14), 1 pd (671-100), 4 IP (671-11-13,15), 13 L (JA 671), 1 pd (673-101), 1 p? (673-100), 1 IP (674-10), 1 p, 4 L (JA 674) [UCLA]. St. Mary: Broadgate, elev. 500 ft, 27 Oct 1966, O.G.W. Berlin and D.C. Watson (JA 643), 4 Ipd (643-32,38-40), 3 lp? (643-34-36), 6 pd (643-104,106-108,112,113), 6 p? (643-30,100-103,105), 2 IP (643-41,42), 3 6,2 %,2P,3 L,51 [UCLA]. St. Thomas: Port Morant Bay, 16 Aug 1967, J.N. Belkin and W.A. Page (JA 729), 3 Ipé (729-10,11,20), 1 pd (729-102), 4 p? (729-100,101,103,104), 7 L, 21 [UCLA]. Windsor Forest, elev. 750 ft, 30 Oct 1966, O.G.W. Berlin and D.C. Watson, 1 Ip? (650-24), 3 pd (650-104,106,107), 1 p? (650-108), 1 IP (650-20), 3 6,29,2L,21(JA 650), 1 Ipd (651-37), 2 lp? (651-3539), 4 pd (651-110-113), 1 p? (651-114), 5 IP (651-31-33,36,38), 1 p, 11 (JA 651), 2 Ipd (652-11,13), 2 pd (652-106,109), 2 p? (652-105, 107), 3 L, 21 (JA 652) [UCLA]. Parish not specified: New Park, June 1929, C.D. Williams, 1 [BM]. Apr 1906, M. Grabham, 1 2; 1944, R.B. Hill, 4 6, 1 9; Jan 1940, W.H.W. Komp, 2 6; R.B. Hill, 1 6, 4 9? [USNM]; W.A. Page, 2 lpdé (771-10,12), 3 pd (771-100,104,107), 3 p? (771-13,102, 105), 1 P, 1 p, 28 L (JA 771), 2 Ip? (772-32,35), 2 pd (772-100,112), 2 p? (772-102,105), 2 d, 5 9,2P,3 p,15 L(JA 772) [UCLA]. Ioliota Group FEMALES. Head: Narrow decumbent scales white. Palpus with apical silvery white scales. Thorax: Inner dorsocentral line narrow, pale golden; median prescutel- lar line broad, white; outer dorsocentral line narrow, pale golden; lateral marginal line narrow, white, nearly continuous from anterior promontory to wingbase. Acros- tichal bristles not developed on disc. Scutellum predominantly dark scaled, light Berlin: Subgenus Howardina 53 scales restricted to median lobe. Ssp with distinct silvery patch; upper stp patch sep- arate, small. Legs: Foretarsus with a few white scales on base of segment 2; midtar- sus with small incomplete basal white rings on segments 1,2; hindtarsus with broader complete basal white rings on segments 1-3 and with segment 5 completely or largely white. Abdomen: Basomedian tergal white markings dingy white or creamy on seg- ments III-VI or V-VI, white on VII. MALES. Palpus with submedian white scales on segments 4,5. Larger claw of fore- leg and midleg with small acute basal tooth and large blunt submedian tooth; smaller Claw with basal tooth only. MALE GENITALIA. Claspette with short broad conical tubercle bearing 1 or rarely 2 apical bristles. Clasper less than 0.5 of sidepiece; with distinct subapical hairs. Aedeagus subquadrate, shorter than proctiger. PUPAE. Cephalothorax: Hair 5-C single or double, at least 2.0 of hair 4-C; hair 7-C usually single; hair 10-C single to triple. Abdomen: All large hairs except 1-IL III, 9-VII,VII usually single or double; hair 2-II laterad of or in line with hair 3; hair 5- IV,V short to moderate, not exceeding middle of tergite following; hair 9-II-VI cau- dad of hair 6; 9-VII longer, stouter and with more branches than hair 6-VII. Paddle: Moderate; apex rounded; marginal spicules indistinct. LARVAE. Head: Hair 1-C unbranched; 4-C moderately long, multiple; 5,6-C both long, single; hairs 11,14-C weakly stellate, 11-C with unequal branches; 15-C double, long, exceeding apex of mental plate. Antenna: Shaft with a few minute sharp spic- ules; hair 1-A single. Thorax: Hair 13-P present. Hairs 0,8,13,14-P, 1,4,5,8,13-T weakly stellate; hair 1-T far mesad of hairs 2,3. Abdomen: Integument glabrous dor- sally, ventral surfaces of segments II-VIII with rows of minute spinelike spicules; hair 2 distinctly removed cephalad but not beyond hair 8, hair 9 not markedly displaced cephalad. Hairs 2,5-I-VI, 9-II-VI, 11-I distinctly stellate, with a few branches, 2,9,11 distinctly heavier than others, hairs 1-I,II, 7-II-VI, 9-I very weakly stellate; hair 8-VII distinctly cephalodorsad of 9-VII. Segment VIII: Comb scales in a single row; indi- vidual scales moderate in length, blunt apically and finely fringed. Siphon: Index less than 3.0; integument without spicules; distinct attached acus present. Pecten extend- ing to about middle of siphon, not reaching hair 1-S; teeth numerous, closely set, with basal denticles. Anal Segment: Saddle body without distinct spicules; caudal margin with short pointed spines preceded by a few rows of spicules; hair 1-X usual- ly single (1-3), near lower end of caudal margin. Ventral brush with 6 pairs of hairs on a small sclerotized boss with distinct grid bars. DISCUSSION. The Ioliota Group of 2 very similar species is most strikingly dif- ferentiated from the other groups of the section by the presence of hair 13-P in the larva, a character not found anywhere else in the section. In adult ornamentation this group is very similar to the Albonotatus Group, differing from it primarily in the presence of light scaling on hindtarsal segment 5. The 2 species have widely disjunct distributions, cozumelensis occurring only on the Yucatan Peninsula and the adjacent Cozumel island and British Honduras, and ioliota at the opposite end of the Caribbean sea in northern Venezuela and Trinidad (fig.3). The 2 species are so similar that they obviously belong to the same phylad. Their pattern of distribution is interpreted here as a relict of a former wide distribu- tion of the group in the central Caribbean area. A similar pattern is found in the Walkeri Group. A larva in a recent collection from a cut bamboo from the province of Guayas (ECU 161) indicates the presence of another member of the Ioliota Group in Ecua- dor. It differs from other members of the group by a 14-16 branched stellate hair 54 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 13-P which is only 3-5 branched in ioliota and double in cozumelensis. The albono- tatus of Levi-Castillo (1952a:77; 1952b:555; 1953:40) is probably this member of the Ioliota Group. Both members of the group were probably originally treehole breeders. They have been reported primarily from artificial containers and ioliota has been found breed- ing in fallen cacao pods. There are no records of this group from bromeliad axils. 12. Aedes (Howardina) ioliota Dyar and Knab Figs.3,8,27,28 1913.Aedes ioliota Dyar and Knab, 1913:76-78. TYPE: Holotype ?, Trinidad, locality not speci- fied, F.W. Urich [USNM, 16000]. Aedes (Howardina) ioliota of Bonne and Bonne-Wepster (1925:371-372); Dyar (1928:233); Ed- wards (1932:155); Lane (1939:100; 1953:719-720); Anduze (1941:14); Horsfall (1955:471); Stone and Knight (1956:219-220); Stone, Knight and Starcke (1959:174); Aitken (1960:4); Suarez and Cova Garcia (1961:17-24); Stone (1963:127); Belkin, Schick and Heinemann (1965: 69); Forattini (1965:401); Cova Garcia, Sutil and Rausseo (1966a:57,283; 1966b:73-74,141, 169,334). Aedes ioliota of Urich (1913:528); Dyar (1918:73). FEMALE. Wing: 3.33 mm. Proboscis: 2.62 mm. Forefemur: 2.45 mm. Abdomen: about 2.98 mm. As described for the group, with following apparently diagnostic fea- tures. Head: Median stripe on vertex narrow, white; light scales on sides and venter straw yellow. Erect scales dark brown. Palpus about 0.16-of proboscis. Thorax (fig. 3): Scales on anterior promontory white. Inner dorsocentral line narrow, creamy, reaching prescutellar space, followed by a moderate, white-scaled median prescutellar line; outer dorsocentral line narrow, golden; lateral marginal line white, narrow, long, extending from side of anterior promontory to wingbase, apparently interrupted on- ly at scutal angle. Antealar area above paratergite with scattered dark scales. Light scales on median scutellar lobe narrow, white; lateral lobes with a few narrow dark scales. Pleural scales as in the section and group. Legs: Forefemur and midfemur pre- dominantly dark anteriorly, a few basal scales creamy, proximal 0.5 of posterior sur- face straw yellow, extending distally as a narrow line, rest dark; hindfemur predomi- nantly straw yellow to 0.5 on both sides, rest dark scaled. Hindtarsal segment 5 com- pletely white scaled. Haltere: Stem pale, knob mostly whitish, a few dark scales dor- sally. Abdomen: Sternites predominantly creamy on II,III; rest with basal creamy and apical dark scales, latter increasing in extent on distal segments. MALE. Wing: 2.79 mm. Proboscis: 2.54 mm. Forefemur: 1.93 mm. Essentially similar to female in coloration. MALE GENITALIA (fig.27). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with 5,6 strong bristles. Sidepiece length about 2.5 median width; basolateral scales white, remainder dark. Claspette with 2,3 small subapical hairs. Clasper about 0.5 of sidepiece, with 1 subapical hair; minute spicules on basal 0.5. Dorsal sclerotization of aedeagus very shallowly emarginate, but without distinct lateral lobes. PUPA (fig.27). Abdomen: 3.35 mm. Trumpet: 0.43 mm; index 4.4-4.6. Paddle: 0.74 mm. Chaetotaxy as given below, hairs moderately to darkly pigmented. Diag-. nostic characters as in the key. Apparently distinguished from cozumelensis by the Berlin: Subgenus Howardina 55 more dendritic hair 1-IJ and longer, ovoid or slightly produced paddle, with length about 1.5 greatest width. Cephalothorax: Mesonotum moderately pigmented between trumpets, lateral areas and metanotum deeply pigmented; trumpets strongly con- trasting. Hair 1(1), 2(2, 1-2), 3(1), 4(2, 1-3), 5(1, 1-2), 6(1), 7(1), 8(3, 2-4), 91), 10(2, 1-3), 111, 1-2), 12(1, 1-3). Abdomen: Tergite I moderately pigmented except for usual membranous areas; segments II and III imbricate, moderately pigmented; rest of segments progressively lighter caudad. Segment I: hair 1(6-12 primary branches, each with many secondary ones), 2(1), 3(1), 4(3, 2-4), 5(2, 2-3), 6(1), 7(1), 9(1). Segment II: hair 0(1), 1(3, strongly dendritic), 2(1, 1-2), 3(1, 1-2), 4(4, 2-5), 5(1), 6(1), 7(1), 9(1). Segment III: hair 0(1), 12, 1-4), 2(1), 31), 4@2, 1-4), 5(1), 6(1), 7(2, 1-4), 8(2, 1-3), 9(1), 10(1,. 1-2), 111), 141). Segment IV: hair 0(1), 1(1, 1-3), 2(1), 3(3, 1-4), 4(1, 1-2), 5(1), 6(1), 7(2, 1-3), 8(2, 1-2), 9(1), 10(2, 1-3), 11(1), 14(1). Segment V: hair 0(1), 1(1, 1-3), 2(1), 31, 1-2), 4(4, 2-5), 51), 6(1), 7(@3, 1- 4), 8(2, 1-3), 9(1), 10(1), 11(1), 14€1, 1-2). Segment VI: hair 0(1), 11, 1-2), 2(1), 3(1, 1-2), 4(2, 1-3), 5(1), 6(1), 701), 8@, 1-3), 9(1), 10€1), 11(1), 14(1). Segment VII: hair 0(1), 1(€1), 2(1), 3(1), 4(€1), 5(1), 6(2, 1-4), 71, 1-2), 8@2, 1-2), 9(3, 2-4), 10(1), 11€1), 14(1). Segment VIII: hair 0(1), 4(1), 9(5, 4-6), 14(1, 1-2). Paddle: Uni- formly lightly pigmented, midrib slightly darker; moderately elongate and oval, sometimes slightly produced; external margin smooth; hair 1-P slender. Male genital lobe extending to 0.47 of paddle, female genital lobe to 0.38. LARVA (fig.28). Head: 0.88 mm. Siphon: 0.86 mm. Anal Saddle: 0.34 mm. Chae- totaxy as given below, hairs moderately to darkly pigmented; in life, larvae are dark. Diagnostic characters as in the key; readily separated from cozumelensis by the api- cally rounded comb scales and hair 1-X single. Head: Width about 1.1 of length; in- tegument strongly pigmented. Mental plate with strong median tooth and 10(9-11) distinct teeth on either side, often a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 413, 12-16), 5(1), 6(1), 7(2, 2-3), 8(1, 1-2), 9(1, 1-2), 10(1), 11(6, 4-7), 12Q, 2-3), 13(1, 1-2), 14(3, 3-7), 15(2, 2-3). Antenna: Length about 0.33 of head; uni- formly strongly pigmented. Hair 1-A about 0.54 from base. Thorax: Long hairs with attenuate apices; long and stellate hairs weakly barbed. Prothorax: hair 0(7, 5-9), 1(3, 2-3), 2(1), 3(4, 3-5), 4(3, 3-5), 5(2, 2-3), 6(1), 7(3, 2-3), 8(6, 5-7), 9(1), 10(1), 11q1, 1-2), 12(1), 13(4, 3-5), 14(20, 18-24). Mesothorax: hair 1(5, 4-6), 2(1), 3(1), 4(1), 5(1), 6(4, 4-5), 7(1), 8(5, 4-6), 9(6, 4-7), 10(1), 11@, 1-2), 12(1), 13(6, 5-9), 14(5, 3-5). Metathorax: hair 1(4, 4-6), 2(1), 3(2, 1-3), 4(6, 3-6), 5(7, 6-8), 6(1), 7(6, 5-7), 8(5, 4-7), 9(5, 3-6), 10(1), 11(1, 1-2), 12(1), 13(6, 5-7). Abdomen: Long and stellate hairs as on thorax. Segment I: 1(4, 4-6), 2(6, 5-7), 3(1), 4(3, 1-4), 5(7, 5-10), 6(3, 2-5), 7(1), 9(4, 3-5), 10(1), 11(6, 6-8), 13(3, 3-5). Segment II: hair 0(1), 1(4, 3- 6), 2(6, 6-7), 3(1), 4(2, 2-4), 5(7, 6-9), 6(3), 7(4, 3-4), 8(1, 1-2), 9(6, 4-7), 101), 11(1), 12(1), 13(5, 3-5). Segment III: hair 0(1), 1(3, 3-5), 2(7, 6-8), 3(1), 4(1), 5(7, 5-10), 6(2), 7(4, 3-5), 8(1), 9(7, 5-7), 10(1), 11(€1), 121), 133, 3-6), 14(1). Segment IV: hair 0(1), 1(4, 1-4), 2(6, 6-7), 3(1), 4(1), 5(8, 6-9), 6(2), 7(4, 3-5), 8(1), 9(6, 5- 8), 10(1), 11(1), 121), 13(4, 3-4), 14(1). Segment V: hair 0(1), 1(2, 1-4), 2(6, 6-7), 3(1), 4(1, 1-2), 5(7, 6-8), 6(2), 7(4, 3-5), 8(1, 1-2), 9(4, 4-7), 10€1), 11€1), 121), 13(3, 2-4), 14(1). Segment VI: hair 0(1), 1(4, 2-5), 2(6, 5-7), 3(1), 4(1), 5(6, 5-8). 6(2, 1-2), 7(3, 2-3), 8(2, 1-2), 9(5, 4-6), 10(1), 11(1), 12(€1), 13(3, 3-4), 141, 1-2), Segment VII: hair 0(1), 1(2, 2-4), 2(6, 5-7), 3(1), 4(1), 5(6, 5-7), 6(2, 2-4), 7(1), 8(4, 3-5), 9(3, 2-4), 10(1), 111), 12(1), 13(2, 1-3), 14(1, 1-3). Segment VIIT: Comb scales in a single row, 16(14-19) in number; individual scale moderate, blunt apically and finely fringed. Hair 0(1), 1(6, 5-7), 2(1), 3(7, 5-8), 41), 5(4, 3-5), 14(1, 1-2). Siphon: Index about 2.3-2.7; darkly pigmented, imbricate. Pecten extending to 0.46; 56 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 teeth 14(12-17); smaller basally, progressively larger distad; individual tooth with a long spine and a basal denticle. Hair 1(3, 2-3), 2(1), 3-5 not visible, 6(1), 7(1), 8(1), 9(1). Anal Segment: Saddle darkly pigmented, imbricate. Gills short, broad, unequal, ventral 0.67 of dorsal. Hair 1(1, 1-3), 2(5, 4-6), 31), 4a(4, 4-6), 4b(4, 3-4), 4c(4, 3- 4), 4d(4, 3-5), 4e(3, 3-4), 4f(3, 1-3). SYSTEMATICS. A. ioliota is extremely similar to cozumelensis from which it is differentiated apparently only by the few constant diagnostic characters mentioned above. There is no indication of any differentiation between the populations from Trinidad and Venezuela. This species is apparently restricted to low and moderate elevations in the Cordillera de la Costa in Venezuela and in Trinidad but may be present in the intermediate area. BIONOMICS. The reported breeding sites of ioliota are quite varied, ranging from rockholes, treeholes, broken and cut bamboo to fallen cacao fruits and artificial con- _tainers of varied size. Females have been taken in biting-landing collections (TR 1132). Attempts at experimental infection with yellow fever virus have yielded nega- tive results (Aitken 1960:4). DISTRIBUTION (fig.3). Trinidad and venesuel (northern) at low to moderate elevations. Material examined: 115 specimens; 15 6, 23 ?, 30 pupae, 47 larvae; 30 in- dividual rearings (12 pupal, 15 larval, 3 incomplete). TRINIDAD. St. Andrew: Cumaca, elev. 250 ft, 22 Oct 1964, rockhole, A. Guerra (TR 783), 1 Ipd (783-104), 2 lp? (783-103,105); elev. 500 ft, 14 Jan 1965, tub, A. Guerra (TR 940), 3 Ip (940-120,121,129), 3 Ip? (940-110,113,125), 1 p? (940-111), 3 IP (940-114,122,123), 12 L; 18 Feb 1965, tin can, A. Guerra, 1 Ip? (1008-21), 1 pd (1008-20), bamboo pot, 2 pd (1013-10,11), 1 p? (1013-13), 5 L (TR 1013) [UCLA]; Cumaca Road, elev. 500 ft, 14 Jan 1965, fallen cacao pod, A. Guerra (TR 938), 4 lp? (938-128-131), 8 L [UCLA] . St. George: Mt. Beck, elev. 1000 ft, 29 Apr 1965, biting collection, A. Guerra, 1 @ (TR 1132) [UCLA] County not specified: F W. Urich, 3 2, type series [USNM no. 16000]. VENEZUELA. Aragua: Choroni, Mar 1939, P.J. Anduze, 1 6 [USNM]. Guamito, 19 June 1927, M.N. Tovar, 1 9; 4 July 1927, M.N. Tovar, 1 2 [USNM]; 1 Dec 1967, cut bamboo, R. Hansell and J.J. Vera (VZ 83), 1 Ipd (83-13), 3 pd (83-101,105,106), 1 p? (83-102), 4 L [UCLA]. Maracay, 4 July 1927, M.N. Tovar, 1 pd [USNM]. Ocumare de la Costa, 12 Aug 1927, M.N. Tovar, 1 ? [USNM]. Colonia Tovar, Capital, Guamitas, Rancho Grande and Uraca (Cova Garcia, Sutil and Rausseo 1966b:73). Carabobo: Las Trincheras (Cova Garcia, Sutil and Rausseo 1966b:73). Fal- con: Cumarebo (Cova Garcia, Sutil and Rausseo 1966b:73). Miranda: Carenero and Charallave (Co- va Garcia, Sutil and Rausseo 1966b:74). Hacienda Los Chaguaramos (Suarez and Cova Garcia 1961: 17; Cova Garcia, Sutil and Rausseo 1966b:74). Los Teques and Petare (Cova Garcia, Sutil and Rausseo 1966b:74). Trujillo: Sabana de Mendoza (Cova Garcia, Sutil and Rausseo 1966b:74). 13. Aedes (Howardina) cozumelensis Diaz Najera Figs.3,8,29 30 1966.Aedes (Howardina) cozumelensis Diaz Najera, 1966a:331-344. TYPE: Holotype 6 with geni- talia slide, San Miguel de Cozumel, Territorio de Quintana Roo, Mexico, 25 Sept 1965, A. Diaz Najera [ISET, 6529]. Aedes (Howardina) cozumelensis of Stone (1967:208). FEMALE. Wing: 3.5 mm. Proboscis: 2.98 mm. Forefemur: 2.28 mm. Abdomen: about 2.5 mm. Very similar to ioliota in color and size, differing apparently only in the following (fig.8). Hindtarsal segment 5 with some apical and posterior dark scales, rest white. Berlin: Subgenus Howardina 37 MALE. Wing: 3.15 mm. Proboscis: 2.45 mm. Forefemur: 2.01 mm. Essentially similar to female in coloration. MALE GENITALIA (fig.29). As figured; diagnostic characters as in the key. Ninth tergite lobe indistinct, with 4,5 strong setae. Sidepiece length about 3.0 median width. Claspette with 1 apical seta and varied number of smaller subapical setae, pro- gressively smaller basally. Clasper about 0.5 of sidepiece, with 1,2 subapical hairs; basal 0.5 with spicules. PUPA (fig.29). Abdomen: 3.78 mm. Trumpet: 0.55 mm; index about 4.1. Paddle: 0.74 mm. Description based on a single skin. Diagnostic characters as in the key. Ap- parently differing from ioliota by hair 1-II with a few simple branches and paddle shorter, length about 1.2 greatest width. Cephalothorax: Mesonotum moderately pigmented between trumpets, lateral areas and metanotum strongly pigmented; trum- pets strongly contrasting. Hair 5-C strong, at least 2.0 as long as 4-C. Abdomen: Ter- gite I strongly pigmented except for usual membranous areas; segments II-IV strong- ly pigmented; rest of segments progressively lighter caudad. All large hairs except 1- II, 9-VII,VIII usually single or double; hair 1-I] not distinctly dendritic and with about 8 branches. Paddle: Uniformly moderately pigmented, midrib slightly darker; slightly broadened in distal half; external margin with submarginal serrations. Male genital lobe extending to about 0.5 of paddle, female genital lobe to 0.36. LARVA (fig.30). Head: 0.76 mm. Siphon: 0.65 mm. Anal Saddle: 0.27 mm. De- scription based on a single specimen. Diagnostic characters as in the key. Hairs mod- erately to strongly pigmented. Readily separated from ioliota by the apically pointed comb scales and hair 1-X double. Head: Hairs and branching as in ioliota. Mental plate with a strong median tooth and 10 distinct teeth on either side, a smaller de- tached tooth at base. Antenna: Slender, moderate; strongly pigmented. Hair 1-A at about 0.56 from base. Thorax: Hair 13-P double. Segment VIII: Comb scales in a sin- gle row, 14 in number; individual scale moderate, pointed apically and fringed. Si- phon: Index about 2.0; imbricate, moderately pigmented, with darker basal ring. Pecten extending to 0.6; teeth 15,16; individual tooth with long spine and a small basal denticle. Anal Segment: Saddle imbricate, moderately pigmented and with darker basal ring. Gills short, broad, unequal, ventral about 0.64 of dorsal. SYSTEMATICS. A. cozumelensis is obviously very closely related to ioliota from which it is separated in all stages apparently only by the few constant features men- tioned in the diagnosis. This species is known from the Yucatan Peninsula and the adjacent Cozumel Island and British Honduras. BIONOMICS. The immature stages of cozumelensis have been found only in arti- ficial containers but it is very likely that the natural breeding sites of this species are similar to those of the closely related ioliota. Adults of cozumelensis have been col- ‘lected at dusk on human bait in British Honduras. DISTRIBUTION (fig.3). Yucatan Peninsula and adjacent islands in Mexico (states of Quintana Roo and Yucatan) and British Honduras. Material examined: 11 speci- mens; 1 6, 8 9, 1 pupa, | larva; no individual rearings. BRITISH HONDURAS. Cayo: Georgeville, elev. 300 ft, 27 July 1967, D.S. Bertram, biting col- lections, 1 ? (BH A369); 1 Aug 1967, D.S. Bertram, 3 ? (BH A339) [UCLA]. Roaring Creek, elev. 70 ft, 27 July 1967, biting collections, D.S. Bertram, 1 + (BH A339); 3 Aug 1967, DS. Bertram, 1 2 (BH 376); 9 Aug 1967, D'S. Bertram, 1 9 (BH 427) [UCLA]. MEXICO. Quintana Roo: Cozumel Island, 25 Sept 1965, A. Diaz Najera, holotype 6 (6529), I paratype 2, 1 p. Yucatan: Tizimin, 8 Dec 1960, M. Munoz, 1 L [ISET]. 58 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Albonotatus Group FEMALES. Head: Narrow decumbent scales white. Palpus with a few apical sil- very white scales. Thorax: Inner dorsocentral line narrow, creamy; median prescutel- lar line broad, white; outer dorsocentral line narrow, pale golden; lateral marginal line narrow, silvery white, extending from side of anterior promontory to wingbase. Acrostichal bristles not developed on disc. Scutellum predominantly dark scaled, light scales restricted to median lobe. Ssp with distinct silvery patch; upper stp patch separate, small. Legs: Foretarsus with a few basal white scales on segment 2; midtar- sus with small incomplete basal white rings on segments 1,2; hindtarsus with broader complete basal white rings on segments 1-3. Abdomen: Basomedian tergal light markings creamy on segments II-VII or III-VII. MALES. Palpus with basomedian white scales on segments 4,5. Larger claw of foreleg and midleg with small acute basal tooth and large blunt submedian tooth; smaller claw simple. MALE GENITALIA. Claspette represented te a basomesal tubercle bearing | api- cal bristle. Clasper subequal to or definitely shorter than di cartons subapical hairs not developed. Aedeagus ovate, subequal to proctiger. PUPAE. Cephalothorax: Hair 5-C double to 4-branched, subequal to hair 4-C and distinctly shorter than distance from its base to that of trumpet; hair 7-C simple to triple; hair 10-C single to 5-branched. Abdomen: All large hairs except 1-II, III, 9-VII, VIII usually single or double; hair 2-II laterad of or in line with hair 3; hair 5-IV,V short, not exceeding apex of tergite following; hair 9-II-VI varied in position; 9-VII longer, stouter and with more branches than 6-VII. Paddle: Very long or moderate; apex rounded or slightly emarginate; marginal spicules distinct. LARVAE. Head: Hair 1-C unbranched; 4-C moderately long, multiple; 5-C single, long; 6-C usually double, strong; none of the ventral hairs stellate; 14-C with a few spikelike branches; 15-C double, long, exceeding apex of mental plate. Antenna: Shaft without distinct spicules; hair 1-A very thin, single. Thorax: Hair 13-P absent. Only hairs 8,13-P distinctly stellate; hair 1-T far mesad of hairs 2,3. Abdomen: Integ- ument glabrous; hair 2 distinctly displaced cephalad but not beyond hair 8, hair 9 not markedly displaced cephalad. Only hairs 2,9-I-VI, 11-I distinctly stellate, with a -few spikelike branches; hair 8-VII distinctly cephalad of 9-VII. Segment VIII: Comb scales in a single or partial double row; individual scales rather short, blunt and faint- ly fringed. Siphon: Index less than 2.5; integument without spicules; distinct free or attached acus present. Pecten not extending beyond middle of siphon; teeth numer- ous, closely set and with basal denticles. Anal Segment: Saddle body without distinct spicules; caudal margin with very short pointed spines preceded by a few spicules; hair 1-X usually double (1-3), near lower end of caudal margin. Ventral brush with 6 pairs of hairs on a moderately or strongly sclerotized boss with distinct grid bars, 1 or 2 hairs usually detached. DISCUSSION. The 2 very similar species of the Albonotatus Group are strikingly differentiated from each other by male genitalic characters, an unusual occurrence in the subgenus. The group is very similar in adult ornamentation to the Ioliota Group, differing from it primarily in having hindtarsal segment 5 completely dark. The lar- vae of the Albonotatus Group lack hair 13-P which is present in the Ioliota Group. Since the loss of hair 13-P is a derived feature it appears that the Albonotatus Group was probably derived from the loliota Group. The Albonotatus Group has a distribution complementary to that of the Busckii Group (fig.3). A. albonotatus has been reported from Hispaniola (Haiti and Domini- Berlin: Subgenus Howardina 59 can Republic) and doubtfully from Guadeloupe, and bahamensis from the Bahamas (Grand Bahama and New Providence). As indicated under albonotatus, a third spe- cies of the group may be present in the Dominican Republic. The natural breeding sites of the group are probably treeholes and broken bam- boo, as has been reported for albonotatus. To date bahamensis has been found only in artificial containers. 14. Aedes (Howardina) albonotatus (Coquillett) i Figs.3,8,31 1905.Gymnometopa albonotata Coquillett, 1905:183. TYPE: Lectotype 6 (141.3) with associated larval and pupal skins, San Francisco Mts., Santo Domingo [Dominican Republic] , 3 Sept 1905, A. Busck [USNM, 8297; selection of Stone and Knight, 1956:213]. Aedes (Howardina) albonotatus of Dyar (1928:233-234); Edwards (1932:155); Gerry (1932:40); Lane (1939:98; 1953:717-719); Horsfall (1955:471); Stone and Knight (1956:213); Stone, Knight and Starcke (1959:173); Belkin, Schick and Heinemann (1965:18); Forattini (1965: 461); Porter (1967:38). Aedes (Howardina) albonotata of Bonne and Bonne-Wepster (1925:369-370). Aedes albonotata of Dyar and Knab (1906b:190,196); Dyar (1906:15); Howard, Dyar and Knab (1912:70; 1917:618,619,834-835); Dyar (1918:73,80); Root (1927:465). Gymnometopa albonotata of Coquillett (1906b:25); Theobald (1907:211; 1910:219); Surcouf and Gonzalez-Rincones (1911:212,213). Haemagogus albonotata of Dyar and Knab (1906a: 166). FEMALE. Wing: 2.9 mm. Proboscis: 2.1 mm. Forefemur: 1.91 mm. Abdomen: about 2.98 mm. As described for the group, with following apparently diagnostic features. Head: Median stripe on vertex narrow, white; light scales on sides and ven- ter straw yellow. Erect scales predominantly dark, a few median ones whitish. Palpus about 0.14 of proboscis. Thorax (fig.8): Scales on anterior promontory white. Inner dorsocentral line narrow, creamy, reaching prescutellar space, followed by a white- scaled median prescutellar line; outer dorsocentral line narrow, golden; lateral mar- ginal line narrow, pure white, extending from sides of anterior promontory to wing- base, apparently interrupted only at scutal angle. Antealar area above paratergite with scattered dark scales. Light scales on median scutellar lobe narrow, white; lateral lobes with a few narrow bronzy scales. Pleural scales as in the section and group. Legs: Forefemur and midfemur predominantly dark, a few basal scales creamy, prox- imal 0.5 of posterior surface mostly straw yellow, extending distally as narrow line, rest dark; hindfemur straw yellow to proximal 0.5 on both sides, rest dark scaled. Hindtarsal segment 3 white scaled in at least basal 0.7. Haltere: Stem pale, knob mostly whitish, a few dark scales dorsally. Abdomen: Basomedian tergal light scales dingy white on II-VII. Sternites straw yellow to segment IV; rest with basal straw yellow and apical dark scales. MALE. Wing: 2.28 mm. Proboscis: 1.93 mm. Forefemur: 1.49 mm. Essentially similar to female in coloration. MALE GENITALIA (fig.31). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with 4 strong bristles. Sidepiece length about 3.0 median width; most tergomesal bristles parallel to longitudinal axis. Clasper narrow, long, subequal to sidepiece; spiniform about 0.2 of clasper. Dorsal sclerotization of aedea- gus slightly emarginate. 60 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 PUPA (fig.31). Abdomen: 2.7 mm. Trumpet: 0.35 mm; index 4.5. Paddle: 0.7 mm. Chaetotaxy as given below. Diagnostic characters as in the key. Readily separated from bahamensis by longer paddle, hair 7-C single and 9-VIII only 3-branched. Ceph- alothorax: Mesonotum and metanotum lightly pigmented; trumpets contrasting. Hair L(1).22(2)s. Shlout- 2) AO 2-3): 5222-3). 601d: 44) BOQ), 9C1), FOC), LIC ye l2ds: 1-2). Abdomen: Tergite I lightly pigmented except for usual membranous areas; pig- mentation of other segments progressively lighter caudad. Segment I: hair 1(9-13 pri- mary branches, each with many secondary ones), 2(1), 3(1), 4(6, 2-8), 5(2, 1-2), 6(1), 7(1), 9€1). Segment II: 0(1), 1(2, 2-3), 21), 31), 4@2, 1-3), 5C1, 1-2), 6(1), 7(1, 1-2), 9(1). Segment III: hair 0(1), 1(1, 1-2), 21), 31), 4@, 2-3), 5(1), 6(1), 7(2, 1-3), 8(1, 1-2), 9(1), 10(1), 11(1), 14(1). Segment IV: hair 0(1), 1@, 1-2), 2(1), 3(1, 1-2), 4(2), 5(2), 6(1), 7(2, 1-2), 8(1), 9(1), 10(1), 11€1), 141). Segment V: hair 01), 11; 1-2), 21), 30); 401, 1-2), 51), 6G): 7@,y 1-2),-8Q; 1-2), 9G); 10), 11(1), 14(1). Segment VI: hair 0(1), 1(1), 2(1), 3(1), 41), 5(1), 6(1), 7(1), 8@, I- 2), 9(1), 10(1), 11(€1), 14(1). Segment VII: hair 0(1), 1€1), 21), 301), 4C1), 5(), 6(1), 71), 8(1), 9(2, 2-3), 10(1), 111), 14(1). Segment VIII: hair 0(1), 4(1), 9(3), 14(1). Paddle: Long; uniformly lightly pigmented, midrib slightly darker; external margin serrated; hair 1-P slender. Male genital lobe extending to 0.55 of paddle, fe- male genital lobe to 0.34. LARVA (Description based on Howard, Dyar and Knab, 1917:853; only frag- ments of two individual rearings seen). Diagnostic characters as in the key. Readily separated from bahamensis by the subequal gills and nature of the comb scales. Head: Width slightly greater than length. Mental plate broadly triangular, with a median tooth and 12 distinct teeth on either side. Antenna: Slender, moderate, uniform; all hairs single. Thorax: Rounded, wider than long. Abdomen: Moderate, anterior seg- ments shorter. Segment VIII: Comb scales in a single row of about 15; individual scale elongate, rounded at tip and with small spinules. Siphon: Short, tapered on outer half, with an index of about 2.0. Pecten extending beyond middle, hair 1-S just within the last tooth; individual tooth with long spine and a small basal denticle. Anal Segment: Saddle as long as wide; caudal margin fringed with spines. Gills about 2.0 length of anal segment; broad and saclike, ventral pair subequal to dorsal. SYSTEMATICS. In spite of the very limited material available and the very close similarity in the ornamentation of. the adults, I consider albonotatus to be undoubt- edly a species distinct from bahamensis. The male genitalia of the 2 species are mark- edly differentiated and the immature stages are also obviously distinct. A. albonotatus has been reported from Haiti, Dominican Republic, Bahamas and Guadeloupe (Stone, Knight and Starcke 1959:173). I have not seen specimens from Haiti but this record may be correct as the type locality of albonotatus is in the Do- minican Republic, also on the island of Hispaniola. The record from the Bahamas is based on a single female from New Providence which in my opinion is bahamensis (see). The male from Guadeloupe collected by Busck [USNM] is unquestionably al- bonotatus and it is possible that this species does occur on this island. However, no additional specimens of albonotatus have been found in extensive surveys on Guade- loupe in recent years and the possibility exists, particularly in view of the extreme similarity of this specimen with males from the type series of albonotatus from the Dominican Republic, that an error occurred in labeling Busck’s material and that this Guadeloupe specimen was actually part of this type series. In addition to the type se- ries I have seen 2 females from the Dominican Republic, near Sanchez [USNM], which differ from the former in the presence of some pale scales on the remigium of the wing. These females are only tentatively identified as albonotatus since they are Berlin: Subgenus Howardina 61 from the northern part of the complex island of Hispaniola and there is a possibility that another species of the group may occur there. The record of albonotatus from Ecuador (Levi-Castillo 1952a:77; 1952b:555; 1953:40) is probably based on a misidentification of an undescribed species of the Ioliota Group (see). BIONOMICS. The type series of albonotatus was reared from larvae and pupae collected in bamboo stumps. DISTRIBUTION (fig.3). Island of Hispaniola (Haiti and Dominican Republic) and doubtfully Guadeloupe. Material examined: 17 specimens; 5 6, 6 9, 4 pupae, 2 larvae; 4 individual rearings (2 pupal, 2 larval). DOMINICAN REPUBLIC. Sanchez, 5 Apr 1949, J.M. Brennan, 2 9? [USNM]. San Francisco Mts, Sept 1905, bamboo, A. Busck, 1 Ipd (141-2), 1 lp? (136-2), 1 pd (136-3), 1 p? (141-7), 1¢ (141), 1 6, 1 9 (136), 1 9 [USNM, UCLA]. ?GUADELOUPE. July 1905, A. Busck, 1 6 [USNM]. HAITI. Mt. Cabrite, 12 Oct 1924, W.A. Hoffman (Root 1927:465). 15. Aedes (Howardina) bahamensis Berlin, n.sp. Figs.3,8 32,33 TYPES: Holotype 6 (BAH 1) with genitalia slide, village of Hunter, Grand Bahama, Bahama Is- lands, 28 Nov 1964, A. Spielman and A.E. Weyer, from colony established from larvae collected in artificial containers [USNM]. Allotype 2 (BAH 1), same data as holotype. Paratypes: 10 6,5 2,9 pupae and 12 larvae (BAH 1), same data as holotype [UCLA]. Aedes (Howardina) albonotatus of Spielman and Weyer (1965:339-343); in part of Stone, Knight and Starcke (1959:173). FEMALE. Wing: 3.5 mm. Proboscis: 2.62 mm. Forefemur: 2.36 mm. Abdomen: about 3.25 mm. Essentially similar to albonotatus but differing by a slightly larger size and in the following (fig.8). Palpus about 0.16 of proboscis. Hindtarsal segment 3 white scaled in barely more than basal 0.5. Basomedian tergal light scales on abdo- men creamy on II-VII. Sternites completely straw yellow up to V, rest with basal straw yellow and apical dark scales. MALE. Wing: 3.06 mm. Proboscis: 2.45 mm. Forefemur: 2.28 mm. Essentially similar to female in coloration. MALE GENITALIA (fig.32). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with 4 strong setae. Sidepiece length about 3.0 median width; basolateral scales white, remainder dark; tergomesal surface clothed with numerous strong, long setae; distal sternomesal margin with threadlike setae with twisted apices. Clasper broad at base, curved mesad on distal 0.7. Dorsal sclerotization of aedeagus deeply emarginate, producing distinct apical lobes. PUPA (fig.32). Abdomen: 4.05 mm. Trumpet: 0.54 mm; index about 4.0. Paddle: 0.78 mm. Diagnostic characters as in the key. Chaetotaxy as given below, hairs dark- ly pigmented. Readily distinguished from albonotatus by the shorter paddle, 7-C usu- ally double and 9-VIII at least 5-branched. Cephalothorax: Mesonotum moderately pigmented between trumpets; sides and metanotum deeper; trumpets strongly con- trasting. Hair 1(1, 1-2), 2(2, 2-4), 3(1), 4(4, 3-6), 5(2, 2-4), 6(1, 1-2), 7(3, 1-3), 8(3, 2-5), 9(1, 1-2), 10(4, 2-5), 11(1), 12(3, 2-5). Abdomen: Tergite I uniformly strongly pigmented except for usual membranous areas; other segments imbricate, strongly pigmented, progressively lighter caudad. Segment I: hair 1(3-10 primary branches, 62 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 each with many secondary ones), 2(1), 3(1), 4(5, 4-6), 5(5, 3-6), 6(2, 1-3), 7(3, 2-4), 9(1), 10(1, present in many), 11(1, present in a few). Segment II: hair 0(1), 1(6, 4- 14), 2(1), 31), 4(6, 4-8), 5(1, 1-2), 6(@, 1-3), 7(3, 2-5), 9(1), 10(2, 1-2, present in a few). Segment III: hair 0(1, 1-2), 1(2, 1-4), 2C), 3(1, 1-2), 4(3, 3-5), 5(1, 1-2), 6(3, 1-4), 7(4, 2-9), 8(3, 2-4), 9(1), 10(2, 2-3), 11€1), 141). Segment IV: hair 0(1), 1€1, 1-2), 2(1), 3(3, 2-4), 4Q, 1-3), 51, 1-2), 6@, 1-3), 7(3, 2-4), 8(3, 2-5), 9(1), 10@, 1-3), 11(1), 12(1, present in one specimen), 14(1). Segment V: hair 0(1, 1-2), 1(2, 1- 3), 21), 3(1, 1-3), 4(3, 2-5), 5(1, 1-2), 6(2, 1-2), 7(3, 2-4), 8(3, 2-4), 9(1), 10(1, 1- 2), 11€1), 14(1). Segment VI: hair O(1, 1-2), 11, 1-3), 2(1), 3(2, 1-2), 4(2, 2-3), 5(1), 6(1), 7(2, 1-3), 8(3, 2-4), 9(1), 101, 1-2), 1101), 141, 1-2). Segment VII: hair Odi, 1-2), 101, 1-3), 2(1), 3@, 1-3), 4€1, 1-2), 51), 6(1, 1-2), 701), 8(2, 1-3), 9(@3, 2- 5), 101), 1101), 14€1). Segment VIII: hair 0(1), 4(1), 909, 5-12), 14(1, 1-2). Paddle: Uniformly lightly pigmented, midrib slightly darker; dorsal surface with fine den- ticles; external margin distinctly serrated and faintly emarginate; hair 1-P slender. Male genital lobe extending to 0.65 of paddle, female genital lobe to 0.44. LARVA (fig.33). Head: 0.92 mm. Siphon: 0.73 mm. Anal Saddle: 0.35 mm. Chae- totaxy as given below, hairs darkly pigmented. Diagnostic characters as given in the key. Readily separated from albonotatus by the unequal gills, pecten teeth termi- nating before middle of siphon and hair 1-S distinctly beyond last pecten tooth. Head: Width about 1.06 of length, moderately pigmented, deeper around mouth- parts. Mental plate with a strong median tooth and usually 12(11-13) distinct teeth on either side, often a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 4Q9, 7- 10), 51), 6(2, 1-2), 7(2, 2-3), 8(1), 9(2, 1-3), 10(1), 1105, 3-6), 12(2, 2-3), 131), 14(2, 2-3), 15(2, 2-3). Antenna: Length about 0.34 of head; moderately pigmented and without spicules; hair 1-A at about 0.5 from base. Thorax: Apices of long hairs attenuate; stellate hairs with a few branches; long and stellate hairs weakly barbed. Prothorax: hair 0(6, 4-6), 1(3, 3-4), 2(1), 3(4, 3-5), 4(3, 2-5), 5(2, 2-3), 6(1), 7(2, 2- 3), 8(7, 5-8), 9(2, 1-2), 10(1), 11(2, 1-3), 12(1), 14(20, 17-29). Mesothorax: hair 1(3, 3-4), 2(1), 3(1), 4€1), 5C1, 1-2), 6(5, 4-5), 7(1), 8(6, 5-9), 9(5, 5-8), 10(1), 111, 1-2), 12(1), 13(6, 3-7), 14(3, 3-4). Metathorax: hair 1(4, 3-5), 2(1), 3(3, 3-5), 43, 2-4), 5(3, 3-5), 6(1), 7(6, 5-7), 8(6, 3-6), 9(5, 4-6), 10(1), 111, 1-2), 121), 13(4, 4+ 7). Abdomen: Stellate hairs as on thorax. Segment I: hair 1(6, 4-7), 2(7, 6-8), 3(1), 4(6, 4-8), 5(6, 4-9), 6(4, 3-5), 7(1), 9(4, 3-5), 10(1), 11(5, 4-7), 13(4, 2-4). Segment II: hair 0(1), 1(3, 2-4), 2(8, 6-8), 3(1), 4(6, 4-6), 5(5, 3-6), 6(4, 3-5), 7(2, 1-3), 8(2), 9(4, 3-5), 10(1), 11€1, 1-2), 12(1), 13(4, 3-5). Segment III: hair 0(1), 1(3, 2-3), 2(7, 6-8), 3(1), 4(2, 1-3), 5(4, 3-7), 6(3, 2-3), 7(5, 4-7), 8(1), 9(6, 5-7), 101, 1-2), 11d], 1-2), 121, 1-2), 13(3, 2-4), 14(1). Segment IV: hair 0(1), 1(2, 2-3), 2(7, 5-8), 31), 4(2, 1-3), 5(5, 4-7), 6(3, 2-4), 7(5, 4-6), 8(1, 1-2), 9(6, 6-9), 10(1, 1-2), 11(1, 1-2), 12(1), 13(2, 2-4), 14(1). Segment V: hair 0(1), 1(3, 2-3), 2(8, 6-9), 3(1), 4(@, 2-3), 5(6, 3-9), 6(2, 2-3), 7(5, 4-5), 8(1, 1-2), 9(7, 6-8), 10(1), 11(2, 1-2), 12(1), 13@, 2- 3), 141, 1-2). Segment VI: hair 0(1, 1-2), 1(3, 2-5), 2(6, 6-8), 3(1), 4(1), 5(4, 3-5), 6(2, 1-2), 7(4, 3-6), 8(2, 2-3), 9(5, 5-7), 10€1), 11(2, 1-3), 12(1), 134, 2-5), 14(@, 2- 3). Segment VII: hair 0(1), 1(4, 3-6), 2(7, 6-8), 3(1), 4C1), 5(4, 3-4), 6(3, 2-4), 71, 1-2), 8(4, 4-5), 9(3, 3-4), 10€1, 1-2), 11€1, 1-2), 12(1), 13(3, 1-4), 14(3, 2-3). Seg- ment VII: Comb scales in a single row, in the form of a wide chevron, 10-18 in num- ber; individual scale moderate; apex blunt and finely fringed. Hair 0(1), 1(4, 2-5), 2(1), 3(7, 6-8), 4(1), 5(4, 2-4), 142, 2-3). Siphon: Index about 2.18-2.4; integument imbricate, moderately pigmented, with darker basal ring. Pecten extending to 0.4, usually 10(6-14) teeth, smaller at base, progressively larger distad; individual tooth with long spine and a basal denticle; hair 1-S distinctly distad of last tooth; acus dis- Berlin: Subgenus Howardina 63 tinct, free or attached. Hair 1(3, 2-4), 2(1), 3-5 not visible, 6(1), 7(1), 8(2, 2-3), 9(1). Anal Segment: Saddle moderately pigmented and imbricate; caudal margin with short spines. Gills short to moderately long, unequal, ventral about 0.5-0.6 of dorsal. Hair 1(2, 1-3), 2(1), 3(4, 4-6), 4a(4, 3-5), 4b(3), 4c(3, 2-4), 4d(3, 3-4), 4e(3, 3-4), 4f(3, 2-4). EGG. Length about 0.67 mm. Width about 0.14 mm; exochorion transparent, with characteristic circular clear refractive bodies; endochorion dark and with large irregular polygonal sculpturing. SYSTEMATICS. I have no hesitation in recognizing bahamensis as a species dis- tinct from the superficially similar albonotatus since the male genitalia are complete- ly different in the 2 forms and the immature stages also appear to be differentiated by constant features mentioned in the diagnosis. In addition to the type series de- rived from a laboratory colony established from material collected on Grand Baha- ma Island and described as albonotatus by Spielman and Weyer (1965:339-343), I have included under bahamensis a single female reared from a treehole on New Prov- idence Island, on which previous records of albonotatus from Bahamas were based. Although this specimen agrees well with the type material, the presence of bahamen- sis on New Providence can be confirmed only by additional material, especially males and immature stages. BIONOMICS. Spielman and Weyer (1965) recorded bahamensis (as albonotatus) from a wide variety of artificial containers similar to those utilized by Aedes aegypti but never in association with the latter. The natural breeding sites of bahamensis are probably treeholes, as indicated for the female from New Providence Island, and pos- sibly also rockholes. According to Spielman and Weyer (loc. cit.) females of baha- mensis are normally autogenous but will occasionally take blood from man or canar- ies. A laboratory colony was readily established and maintained; eggs produced by autogenous females hatched in tap water after storage for 1-2 weeks at a relative hu- midity of 76% and a temperature of 76°F. DISTRIBUTION (fig.3). Bahama Islands. Material examined: 45 specimens; 13 d, 8 2,9 pupae, 15 larvae; no individual rearings. BAHAMA ISLANDS. Grand Bahama: village of Hunter, artificial containers, A. Spielman (BAH- 1, type series), 11 6,6 2,9 p, 12 L [USNM, UCLA]; 19 July 1964, tire, A. Spielman, 1 6 genitalia (GB 71-1), 1 L; 1 Dec 1964, tire. A. Spielman, 1 6 (GB 81-52A), 1 9 (GB 81-85A) [USNM] . New Providence: 1915, treehole, H.G. Dyar, 1 9 [USNM]. Busckii Group FEMALE. Head: Narrow decumbent scales pale golden. Palpus completely dark scaled. Thorax: Inner dorsocentral line faint, narrow, golden; median prescutellar line narrow, golden; outer dorsocentral line faint, narrow, golden; lateral marginal line narrow, silvery, interrupted at scutal angle and sometimes caudad of it. Acros- tichal bristles present on disc. Scutellum with some light scales on lateral lobes. Ssp with distinct silvery patch; upper stp patch small, separate. Legs: Foretarsus com- pletely dark; midtarsus with a few basal white scales on segments 1,2; hindtarsus with short to moderate basal white rings on segments 1-3. Abdomen: Basomedian tergal light markings evident from segment V distad. MALE. Palpus entirely dark. Larger claw of foreleg and midleg with submedian tooth only; smaller claw without either submedian or basal tooth. MALE GENITALIA. Claspette with a distinct outstanding apical fingerlike cylin- 64 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 — drical process more than twice as long as wide and bearing a short apical bristle. Clasper about 0.6 of sidepiece; subapical hairs absent. Aedeagus ovate; apex deeply - emarginate, with distinct lateral lobes; subequal in length to proctiger. PUPA. Cephalothorax: Hair 5-C double to 4-branched, short but longer than hair 4-C; hair 7-C double to 4-branched; hair 10-C with 2-5 branches. Abdomen: All large hairs except 1-II, III, 9-VII, VIII usually single or double; hair 2-II laterad of or in line with hair 3; hair 5-IV,V short, usually not reaching apex of tergite following; hair 9- II-VI usually caudad of level of hair 6; 9-VII longer, stouter and with more branches than hair 6-VII; hair 4-VIII usually single. Paddle: Short; apex rounded; marginal spicules distinct. LARVA. Head: 1-C unbranched; 4-C moderately long, multiple; 5-C single, long; 6-C usually double or triple; hairs 11,14-C weakly stellate, 11-C with unequal branches; 15-C usually double, long, exceeding apex of mental plate. Antenna: Shaft with a few scattered weak sharp spicules; hair 1-A single. Thorax: Hair 13-P absent. Hairs 0,8,14-P, 1,13,14-M, 1,4,5,8,13-T all moderately stellate and moderately pigmented; hairs 1-M,T far mesad of hairs 2,3. Abdomen: Integument glabrous; hairs 2 and 9 far removed cephalad but not beyond hair 8. Hairs 1,2,5,9,13-I-VI, 11-1, 7-III-VI all moderately stellate, usually stronger than on thorax; hair 8-VII cephalodorsad of 9- VII. Segment VIII: Comb scales varied from 1 more-or-less regular row to 2-4 dis- tinct rows; individual scales moderately long and broad, blunt apically and fringed. Siphon: Index less than 3.0; integument without spicules; small separate acus pres- ent. Pecten variable, sometimes extending to near apex; teeth always few and widely spaced, short, proximal with basal denticles. Anal Segment: Saddle body without distinct spicules; caudal margin with relatively short fringed spines; hair 1-X usually less than 5-branches, near lower end of caudal margin. Ventral brush with 5 pairs of hairs on a moderately sclerotized boss with short median grid bars. DISCUSSION. Only 1 species is recognized at this time in the Busckii Group al- though it is evident, as indicated under the species, that there are marked differences among the populations from different islands. The Busckii Group is immediately separated from all the other groups in the sub- genus by the presence of acrostichal bristles on the disc of the mesonotum. The or- namentation of the adults is in general intermediate between the Walkeri Group and the Ioliota-Albonotatus Groups. The larvae of some populations show another prim- itive feature in the presence of several rows of comb scales, a condition found nor- mally only in the Fulvithorax Group. Since the distributions of these 2 groups are now adjacent and may have overlapped in the past, the comb patch in busckii may have been acquired through introgression from the Fulvithorax Group. This does not seem likely because this feature is found only in the northern populations, farthest removed from Fulvithorax Section and because there is no indication of any other introgression between the 2 groups. The Busckii Group is definitely known only from the Lesser Antilles from Sint Eustatius southward to Grenada (fig.3). This distribution is complementary to the Ioliota and Albonotatus groups except for a possible overlap with the latter in Gua- deloupe. The Busckii Group has the widest range of breeding sites in the subgenus, being found in treeholes, broken bamboo, rockholes, fallen fruits and leaves, as well as in leaf axils of bromeliads and aroids, and in flower bracts of heliconias. Berlin: Subgenus Howardina 65 16. Aedes (Howardina) busckii (Coquillett) Figs.3,8,34,35 1906.Stegomyia busckii Coquillett, 1906:60-61. TYPE: Lectotype d, Cacao plantation, San Do- mingo [Dominica], 28 July 1905, A. Busck [USNM, 9139; selection of Stone and Knight $956: 215). 1919.Aedes thaxteri Dyar and Knab, 1919:8-9. TYPE: Holotype 6, Grand Etang, Grenada, Nov 1912, R. Thaxter [USNM, 21704] . Synonymy with busckii by Dyar and Knab (1919:8). 1941.Aedes tracei Senevet and Quievreux, 1941:251-258. TYPE: Holotype 6 (M 2163) with asso- ciated pupal skin and genitalia mount, near Piscine Colson, Martinique, 14 Aug 1939, G. Senevet and L. Quievreux [NE; see Belkin 1968]. Synonymy with busckii by Belkin, Schick and Heinemann (1965:33). Aedes (Howardina) busckii of Bonne and Bonne-Wepster (1925:372-373); Dyar (1928:234-235); Edwards (1932:155); Lane (1939:99); Floch and Abonnenc (1945:14-16,42,45,47); Lane (1953:722-724); Stone, Knight and Starcke (1959:173); Belkin, Schick and Heinemann (1965: 17,33); Porter (1967:38). Aedes busckii of Dyar and Knab (1906b:189,192); Howard, Dyar and Knab (1917:860-862); Dyar (1918:73,80); Senevet (1938:189); van der Kuyp (1954:39,64,68,70,72,95,96); Stone and Knight (1956:215). Gymnometopa busckii of Coquillett (1906b:25); Theobald (1907:211; 1910:219); Surcouf and _ Gonzalez-Rincones (1911:212,213). | Haemagogus busckii of Coquillett (1905 :183); Dyar and Knab (1906a: 166). Aedes (Howardina) thaxteri of Bonne and Bonne-Wepster (1925:276-277); Lane (1939:99). Aedes (Soperia) tracei of Lane (1953:703-705); Stone, Knight and Starcke (1959:173); Forattini (1965 :460). | FEMALE. Wing: 3.24 mm. Proboscis: 2.19 mm. Forefemur: 2.19 mm. Abdomen: about 3.0 mm. As described for the group and with following additional features. Head: Light scales on median stripe pale golden; creamy on sides and venter. Thorax (fig.8): Light patch on anterior promontory silvery. All mesonotal light lines, except lateral marginal, narrow, golden. Antealar area above paratergite with scattered dark scales. Lateral scutellar lobes with a few narrow golden and dark scales. Legs: Fore- femur and midfemur predominantly dark anteriorly, a few creamy scales at base, posterior surface with a lateral and a median longitudinal creamy line from base to apex on forefemur and midfemur respectively, rest dark; hindfemur predominantly creamy to 0.6, lateral margin and distal area dark. Haltere: Stem pale, knob predom- inantly dark, fringe scales whitish. Abdomen: Sternites completely creamy on II-IV; rest of segments with basal creamy and apical dark scales. MALE. Wing: 2.54 mm. Proboscis: 2.19 mm. Forefemur: 1.84 mm. Essentially similar to female in color. MALE GENITALIA (fig.34). As figured; diagnostic features as described for group. Ninth tergite lobe indistinct, with 2,3 strong bristles. Sidepiece elongate, length about 3.5 median width. PUPA (fig.34). Abdomen: 3.35 mm. Trumpet: 0.43 mm; index about 4.0. Paddle: 0.7 mm. Chaetotaxy as given below, hairs moderately pigmented. Diagnostic charac- ters as in the key. Cephalothorax: Mesonotum lightly to moderately pigmented be- tween trumpets; lateral areas and metanotum deeper; trumpets strongly contrasting. Hair 1(1), 21, 1-2), 3(1), 403, 2-4), 5@, 2-4), 6(1, 1-2), 7@2, 2-4), 8@, 1-3), 9(1), 10(3, 2-5), 11(1), 12(2, 1-3). Abdomen: Tergite I moderately pigmented and imbri- cate except for usual membranous areas; pigmentation progressively lighter on distal 66 | Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 segments. Segment I: hair 1(9-14 primary branches, each with many secondary ones), 2(1, 1-2), 3(1), 4(4, 3-5), 5(3, 2-6), 6(1, 1-2), 7@, 1-3), 91), 111, present in a few). Segment II: hair 0(1), 1(4, 3-7), 21, 1-2), 31, 1-2), 4(4, 2-5), 5C1, 1-3), 61, 1-2), 7(2, 1-3), 8(1, 1-3, present in many), 9(1), 11(1, present in some). Segment III: hair O(1), 1(2, 1-3), 21, 1-3), 31, 1-2), 4(3, 2-4), 51, 1-4), 6(1, 1-2), 7(3, 2-4), 8(2, 2-3), 9(1), 10(2, 1-2), 11(1), 14(1). Segment IV: hair 0(1), 1(2, 1-3), 2(1), 3(3, 2-4), 4Q, 2-3), 5(2, 1-3), 6(1), 702, 2-4), 8(2, 1-3), 9(1), 10(2, 1-3), 11(€1), 14(1). Segment V: hair O(1), 11, 1-2), 2(1), 3@, 1-3), 4(3, 3-5), 5(1, 1-3), 6(1), 7(3, 2-3), 8(2, 1-4), 9(1), 10(1), 11(1), 14(1). Segment VI: hair 0(1), 1(1, 1-2), 2(1), 3(@2, 1-3), 4(@2, 2-3), S(1, 1-2), 6(1), 7(2, 1-3), 8(2, 1-3), 9(1), 10€1), 11(1), 14(1). Segment VII: hair 0(1), Id, 1-2), 2(1), 3(1), 4@, 1-2), 5(1, 1-2), 6(1, 1-3), 7€1), 8(1, 1-2), 9(5, 3-7), 10C1, 1-2), 11(1), 14(1). Segment VIII: hair 0(1), 4(1, 1-2), 9(7, 5-11), 141, 1-2). Paddle: Uniformly lightly pigmented, midrib slightly darker; length subequal to or slightly greater than median width; dorsal surface with minute denticles; apex rounded, ex- ternal margin serrated; hair 1-P slender. Male genital lobe extending to about 0.45 of paddle, female genital lobe to 0.36. : LARVA (fig.35). Head: 0.73 mm. Siphon: 0.59 mm. Anal Saddle: 0.27 mm. Diag- nostic characters as in the key. Readily separated from other members of the section by the presence of only 5 pairs of hairs in ventral brush. Chaetotaxy as given below, hairs moderately to darkly pigmented. Head: Width about 1.08 of length; integument moderately pigmented, deeper around mouthparts. Mental plate with a strong medi- an tooth and usually 12(10-13) distinct teeth on either side, often a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 4€11, 10-12), 5(1), 6(2, 2-3), 7(3, 2-5), 8(1), 91, 1-3), 10(1), 11(5, 4-10), 12(2, 2-3), 13(1), 14(3, 2-6), 15(2, 2-4). Antenna: Length about 0.3 of head; uniformly moderately pigmented, with a few scattered spicules. Hair 1-A at about 0.6 from base. Thorax: Apices of long ha?rs attenuate; stellate hairs with a few to a moderate number of branches; long and stellate hairs weakly barbed. Prothorax: hair 0(7, 6-9), 1(4, 2-5), 2(1), 3(5, 4-8), 4(4, 4-5), 5(2, 2-3), 6(1),.7(2), 8(7, 5-12), 9(2, 1-3), 10(1), 11(3, 2-3), 12(1), 14(20, 19-29). Mesothorax: hair 1(8, 4-12), 2(1), 3(1), 4(1), 51), 665, 3-5), 7(1), 865, 5-7), 9(6, 4-8), 10(1), 11(1, 1-3), 12(1), 13¢5, 4-8), 14(7, 3-10). Metathorax: hair 1(6, 3-12), 2(1), 3(2, 1-3), 4(7, 4- 10), 5(7, 5-10), 6(1), 7(5, 5-7), 8(5, 4-6), 9(4, 4-6), 10(1), 11(2, 1-3), 12(1), 1366, 6-13). Abdomen: Integument and stellate hairs as on thorax. Segment I: hair 1(7, 4- 11), 2(10, 7-13), 3(1), 4(2, 1-3), 5(7, 7-12), 6(5, 3-6), 7(1), 9(4, 3-5), 10(1), 1109, 7-12), 13(5, 3-6). Segment II: hair 0(1), 1(8, 5-12), 2(12, 7-12), 3(1), 4(2, 1-2), 5(9, 7-11), 6(4, 3-6), 7(4, 2-4), 8(1), 9(5, 4-7), 10(1), 11€1), 12(1), 13(7, 4-12). Segment Hit: hair 0(1), 1(7, 5-12), 210, 8-10), 3(1), 4(1), 5(8, 8-14), 6(2, 2-3), 7(5, 4-7), 8(1), 9(9, 6-10), 10(1), 11€1), 12(1), 13(8, 4-12), 14(1, 1-2). Segment IV: hair 0(1), 1(10, 7-10), 2(10, 7-10), 3(1), 4(1), 59, 8-13), 6(2), 7(5, 4-7), 8(1, 1-2), 9(8, 8-10), 10(1), 11(1), 12(1), 13(8, 5-12), 14(1). Segment V: hair 0(1), 1(5, 5-12), 2(10, 7-10), 3(1), 4(1, 1-2), 5(8, 8-12), 6(2), 7(5, 4-8), 8(1), 9(9, 7-10), 10(1), 11(1), 12(1), 13(7, 5- 11), 14€1, 1-2). Segment VI: hair 0(1), 1(9, 5-12), 2(8, 5-10), 3(1), 4(1), 5(9, 7-13), 6(2), 7(3, 3-5), 8(2, 1-3), 9(7, 6-10), 10(1), 11(1), 12(1), 13(5, 3-10), 14(2, 1-2). Segment VII: hair 0(1), 109, 4-12), 2(7, 5-10), 3(1), 4(1), 5(7, 5-11), 63, 3-5), 7(1, 1-2), 8(6, 5-8), 9(3, 2-4), 10(1), 11(1), 12(1), 13(3, 3-7), 14(1). Segment VIIT: Comb scales varied from a single regular row to a distinct patch of 3,4 rows; individual scale long, blunt apically and fringed. Hair 0(1), 1(10, 6-12), 2(1), 3(7, 6-7), 4(1), 5(6, 3-10), 14(1, 1-2). Siphon: Index about 2.2-2.8; integument moderately pigmen- ted, with darker basal ring. Pecten teeth few (6-10), widely spaced; frequently teeth terminate before hair 1-S, occasionally 1,2 teeth extend beyond it; individual tooth Berlin: Subgenus Howardina 67 with a long spine and small basal denticle; acus distinct and separate or indistinct. Hair 1(2, 2-4), 2(1), 3-5 not visible, 6(1), 7(1), 8(1), 9(1). Anal Segment: Saddle moderately pigmented; caudal margin with moderately large flat spines; individual spine with lateral spinules. Gills short to moderate, broad, unequal, ventral about 0.6 of dorsal. Hair 1(2, 2-4), 2(4, 4-6), 3(1), 4a(4, 3-6), 4b(3, 3-4), 4c(4, 3-4), 4d(4, 4-5), 4e(2, 2-3). | SYSTEMATICS. A. busckii in the present interpretation is undoubtedly a complex of closely related and more or less strongly differentiated forms, some of which may be geographically or ecologically isolated species. It is impossible to resolve this com- plex at this time because of the bewildering array of differentiation and variation in ornamentation of the adults and larval characters which is apparently associated with both isolation on numerous islands and the utilization of a wide variety of breeding sites (see Bionomics below). As in the case of the 7ripteroides caledonicus complex in the South Pacific (Belkin 1955), the situation may be further complicated through recent mixing of originally isolated insular populations by human transport. For the present, the currently accepted synonymy of thaxteri with busckii is re- tained although there are rather obvious differences in ornamentation of the adults as pointed out in the original description of the former. The synonymy of tracei, proposed by Belkin, Schick and Heinemann (1965:33), appears to be well founded since the populations from Martinique are very similar to those from Dominica. The above diagnosis of the complex is based primarily on topotypic associated ma- terial of all stages of busckii from Dominica. In the adults, the most striking differ- ences in the various populations are: (1) in the nature of the lateral mesonotal silvery line which may be nearly continuous from the anterior promontory to the wing root or variously broken into segments, (2) in the extent of the basal white ring on hind- tarsal segment 3 which varies from less than 0.4 to almost 0.75 of the segment length, and (3) the presence of some white scales on additional tarsal segments. No striking differences have been noted in the pupae but the larvae of different populations show remarkable differentiation from the typical population in: (1) the number and ar- rangement of scales in the comb from a few scales in a single even row to a large number in a patch of 3 or 4 rows, and (2) the distribution of the pecten which may end before the siphonal tuft or may reach nearly to the tip of the siphon. The northern populations from Sint Eustatius, St. Kitts, Montserrat and Guade- loupe differ from all the others in the more extensive white ring on hindtarsal seg- ment 3, the larval comb in a large patch and the pecten ending before the siphonal hair tuft. The male genitalia of these populations are also apparently distinct from the others in the large number of setae on the tergal surface of the sidepiece and in a broad aedeagus with distinct apicolateral lobes. Although these populations probably represent a distinct species, I prefer not to name or describe it formally without addi- tional material and a thorough study of the entire complex which is beyond the scope of the present revision. The busckii complex is apparently restricted to the main chain of the Lesser An- tilles from Sint Eustatius southward to Grenada. I have seen material only from the above-mentioned islands and from St. Kitts, Montserrat, Guadeloupe, Dominica, Martinique and St. Lucia. No material of the complex was found in a recent survey of Nevis and St. Vincent but it is likely that populations will be found there as well as on the smaller intermediate islands of the chain. The report of busckii from Jamai- ca by Hill and Hill (1948:43) is probably based on an erroneous identification of a member of the Aurites Group and its record from the Dominican Republic is due to the error in labeling the type material San Domingo instead of Dominica (see How- 68 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 ard, Dyar and Knab 1917:860). BIONOMICS. Members of the busckii complex have been found breeding in the entire range of habitats utilized by the subgenus, in rockholes, treeholes, broken and cut bamboo, fallen leaves, fruits and pods, in the leaf axils of bromeliads and aroids, and in the flower bracts of heliconias. The immature stagés are also commonly found in a variety of artificial containers. One male (FWI 245) from Guadeloupe is re- ported from a ground pool but this may be a contamination or an error in labeling. Females are frequently taken in biting-landing collections and some were obtained on chicken bait. Apparently some adult populations may reach high levels and be- come serious pests locally. DISTRIBUTION (fig.3). Main chain of the Lesser Antilles from Sint Eustatius to Grenada; reported from low (40 ft) to moderate (2600 ft) elevations. Material exam- ined: 1269 specimens; 253 6, 273 9, 290 pupae, 453 larvae; 175 individual rearings (19 pupal, 131 larval, 25 incomplete). DOMINICA. St. Andrew: Calibishie, 23 June 1965, colocasia leaf axil, R. Martinez and A. Guer- ra, 1 p, 21 (DOM 137), 2 lpd (138-10,11), 1 lp? (138-13), 1 P (DOM 138) [UCLA]. St. David: Castle Bruce, 19 Apr 1959, bamboo, R.H. Darsie, 1 Ipd (76-14), 2 pd (76-15A,16), 1 d (76-10), 1 lp? (76-6), 4 2 (76-2,4A,7,11), 1 IP, 1 P [USNM]. Grand Fond, elev. 300 ft, 18 June 1965, biting collection, R. Martinez and A. Guerra, 1 9? (DOM 116) [UCLA]. Terre Ferme, elev. 1800 ft, 22 June 1965, heliconia bract, R. Martinez and A. Guerra (DOM 131), 1 lpd (131-12), 1 Ip? (131-11), 6 6,12 p,2L [UCLA]. St. George: L’Etang, elev. 2600 ft, 8 June 1965, bromeliads, T.H.G. Ait- ken, R. Martinez and A. Guerra, 1 p (DOM 53) [UCLA]. Trafalgar Falls, elev. 1700 ft, 9 June 1965, biting collection, R. Martinez and A. Guerra, 4 ? (DOM 58); heliconia bract (DOM 59), 2 lp? (59-10,11), 1 P, 1 p [UCLA]. St. Joseph: Batati Estate, elev. 40 ft, 14 June 1965, treehole, R. Mar- tinez and A. Guerra (DOM 82), 1 Ip? (82-20) [UCLA]. Clarke Hall Estate, 12 Nov 1964, P.J. Span- gler, 1 9; 11 Mar 1964, D.F. Bray, 2 9; 21-31 Jan 1965, W.W. Wirth, 3 ?; 24 Jan 1965, W.W. Wirth, 2 2;3 May 1966, R.J. Gagne, 1 9; 11 May 1966, G. Steyskal, 1 9; 19 May 1966, G. Steyskal, 1 9; 24 May 1966, G. Steyskal, 3 2; 1 June 1966, G. Steyskal, 1 9; 3 June 1966, G. Steyskal, 1 2; 10 June 1966, G. Steyskal, 6 9; 11 June 1966, G. Steyskal, 2 9; 12 June 1966, G. Steyskal, 1 9; 10 Oct 1966, A.B. Gurney, 1 9 [USNM]. St. John: Milton Estate, elev. 1000 ft, 24 June 1965, heli- conia bract, A. Guerra (DOM 159), 2 Ips (159-10,13), 1 pd (159-101), 2 P, 4 p, 14 L [UCLA]. St. Luke: Point Michele, Champigny Estate, elev. 600-700 ft, 6 June 1965, treehole, T.H.G. Aitken, R. Martinez and A. Guerra, 2 Ipd (15-20,22), 1 Ip? (15-23), 1 P, 1 p, 1 1(DOM 15), 1 lpd (19-10), 4 6,8 p, 7 L(DOM 19), 4 L (DOM 20), fallen leaf, 1 Ip? (17-10), 2 p, 6 L, 21(DOM 17), steel drum, 3 6, 1 2? (DOM 21), heliconia bract, 4d, 1 9? (DOM 23) [UCLA]. St. Mark: Tete Morne, elev. 1400 ft, 10 June 1965, heliconia bract, R. Martinez and A. Guerra (DOM 68), 2 Ipd (68-12,14), 2 Ip? (68-11,13), 4 6, 8 p, 4 L [UCLA]. St. Paul: Antrim Valley, 7 June 1965, treehole, R. Mar- tinez and A. Guerra, 4 9, 4 p, 4 L (DOM 35) [UCLA]. Mount Joy, elev. 1000 ft, 7 June 1965, treehole, R. Martinez and A. Guerra (DOM 41), 1 Ipd (41-15), 2 Ip? (41-10,11), 1 P, 11 L [UCLA]. Pont Casse, 23 Nov 1964, P.J. Spangler, 1 6 [USNM]. Parish not specified: July 1905, in sugar boiler full of vegetation, 1 6, 1 2 (78), type series [USNM no. 9139]. GRENADA. St. Andrew: Balthazar Estate, elev. 450 ft, 15 Oct 1963, bamboo, R. Martinez, 1 Ipd (43-125), 1 Ipd (44-136), 1 Ip? (44-131) [UCLA]. Mt. St. Catherine, elev. 2000 ft, 25 Oct 1963, bamboo, R. Martinez, 2 L (GR 82), heliconia bract, 2 ? (GR 85) [UCLA]. St. David: Minor- ca Estate, elev. 700 ft, 24 Oct 1963, colocasia leaf axil, R. Martinez, 3 L (GR 76); 1 Nov 1963, fal- len cacao pod, R. Martinez (GR 111), 1 lpd (111-18) [UCLA]. St. George: Annandale, elev. 700 ft, 25 Aug 1929, leaf axil, F.M. Root, 18 6, 24 9? (LAR 40), 5 2? (LAR 40A); 28 Oct 1963, bamboo, R. Martinez, 1 L (GR 95), rockhole (GR 96), 1 Ip (96-156) [UCLA]. Grand Etang, 2 July 1929, F.M. Root, 20 6, 40 9 (GRR 11); 26 Aug 1929, leaf axil, F.M. Root, 21 6,4 ? (LAR 42); 17 Oct 1963, heliconia bract, R. Martinez (GR 54), 1 Ip? (54-132), bamboo (GR 59), 1 Ipd (59-133), fallen palm leaf, 9 6,3 2 (GR 60), heliconia bract (GR 61), 1 Ipd (61-127), 1 Ip? (61-139); 18 Oct 1963, biting collection, R. Martinez, 1 9 (GR 62); 23 Oct 1963, heliconia bract, R. Martinez (GR 67), 1 lpdé (67-140), 1 6 [UCLA]. St. Mark: Concord, F.M. Root, 27 d, 15 2? (GRR 80) [UCLA]. Berlin: Subgenus Howardina 69 Parish not specified: FM. Root, 11 6, 29 ? (GRR 40) [UCLA]. GUADELOUPE. Basse-Terre: Capesterre, 76 L, Matouba, 72 L; Petit-Bourg, 17 L; Pointe-Noire, 12 L (Floch and Abonnenc 1945:16). Route de Traversee, km 4-6, elev. 200 ft, 18 Sept 1965, ground pool, P. Fauran, 1 6 (FWI-245) [UCLA]. St. Claude, Bains Jaunes, elev. 900 ft, 31 Mar 1965, heliconia bract, P. Fauran, 2 Ipd (211-12,13), 5 lp? (211-10,11,15-17), 2 IP (211-14,18), 4 Ipd (212-11,13,15,16), 2 Ip? (212-10,14), 1 IP (212-12) [UCLA]. Soufriere, 111 L; Trois- Rivieres, 51 L (Floch and Abonnenc 1945:16). Vernou, elev. 300 ft, 20 July 1964, treehole, P. Fauran (FWI 182), 4 lpd (182-101,102,105,107), 6 lp? (182-103,106,108,110,112,113), 1 IP (182- 111), 2 L [UCLA]. Locality not specified, elev. 3000 ft, July 1905, wild canna, A. Busck, 1 Ipd (80-10), 2 ? (80-3,7), 36,49 [USNM]. MARTINIQUE. Fort de France, 1929, bamboo, F.M. Root, 2 6 (LAR 23), bromeliads, 1 9 (LAR 28A) [UCLA]; Route de la Trace, elev. 680 ft, 16 July 1965, heliconia bract, P. Fauran (FWI-M2), 2 Ipd (M2-10,11), 1 Ip? (M2-14), 3 pd (M2-15,101,102), 2 p? (M2-12,100), 1 IP (M2- 13),3 p, 11 [UCLA]. Mt. Pelee, July 1905, A. Busck, 2 9? [USNM]. Piscine Colson, 14 Aug 1939 (Senevet and Quievreux 1941:263). MONTSERRAT. St. Antony: Chance Mountain, elev. 2600 ft, 5 Oct 1966, treehole, T.H.G. Aitken, R. Martinez and A. Guerra, 2 P, 10 L(MNT 25), 2 lpd (28-10,11), 1 lp? (28-13), 1 pd (28- 100), 2 IP (28-12,14), 19 L (MNT 28), 1 Ipd (29-10), 2 lp? (29-11,12), 3 L (MNT 29), heliconia bract (MNT 27), 4 Ipd (27-12,13,15,16), 2 lp? (27-10,14), 1 pd (27-100), 1 IP (27-17), 5 L[UCLA]. Parsons Estate, elev. 500 ft, 13 Oct 1966, tin can, R. Martinez and A. Guerra (MNT 72), 1 IP (72- 10),9 L [UCLA]. Soufriere, elev. 1000 ft, 12 Oct 1966, bamboo, R. Martinez and A. Guerra, 1 IP (66-10), 1 p, 5 L(MNT 66), 5 L (MNT 67) [UCLA]; Lower Gages, 1250 ft, 6 Oct 1966, treehole, R. Martinez and A. Guerra, 8 L (MNT 36) [UCLA] ; Upper Gages, 1750 ft, 6 Oct 1966, heliconia bract, T.H.G. Aitken, R. Martinez and A. Guerra (MNT 38), 1 Ipdé (38-10), 1 pd (38-100), 1 p? (38-101), 5 L [UCLA]. Waterworks Estate, elev. 550 ft, 18 Oct 1966, treehole, R. Martinez and A. Guerra (MNT 118), 1 p? (118-100), 1 IP (118-20) [UCLA]. St. Peters: Plymouth, Belham Bridge, elev. 250 ft, 4 Oct 1966, tin can, T.H.G. Aitken, R. Martinez and A. Guerra (MNT 20), 2 Ipd (20-11,12), 1 Ip? (20-10), 1 pd (20-100), 1 p? (20-101), treehole (MNT 22), 1 lpd (22-14), 3 Ip? (22-10-12), 1 IP (22-13), 9 6, 3 9, 12 p [UCLA]. SINT EUSTATIUS. 7 Oct 1947, E. van der Kuyp, 2 6, 1 9 [USNM]. ST. KITTS. Christ Church: Old Road Town, elev. 1200 ft, 5 Sept 1966, heliconia bract, A. Guerra, 1 6, 2 9; 5 Sept 1966, R. Martinez and A. Guerra, 1 p? (93-101), 2 6, 1 9 (KIT 93), 1 6, 4? (KIT 94) [UCLA]. St. John: Dieppe Bay Town, slopes of Mount Misery, elev. 2500 ft, 17 Aug 1966, treehole, T.H.G. Aitken, R. Martinez and A. Guerra (KIT 25), 1 lp? (25-10), 1 IP (25-11), Oc, 2 %,3 pi. (UCLA): ST. LUCIA. Anse la Raye: Elev. 300 ft, 15 July 1964, treehole, R. Martinez and A. Guerra, 1 L (LU 30); 30 July 1964, fallen fruit, A. Guerra, 1 lp? (125-20), 1 p? (125-100), 1 lp (126-20), 4 L(LU 126), treehole (LU 128), 7 Ip? (128-10-16), 1 IP (128-17), 26 6, 10 9, 19 P, 11 L [UCLA]; Derriere Dos, 31 July 1964, fallen cacao pod, A. Guerra (LU 131), 1 lpd (131-24), 5 1p? (131-20- 23,25), 4 6, 12 ?, 8 p [UCLA]. Barre de L’Isle, elev. 800-900 ft, 14 July 1964, treehole, T.H.G. Aitken, R. Martinez and A. Guerra, 3 lpd (14-14-16), 6 lp? (14-10-13,18,19), 8 p, 7 L, 3 1(LU 14), 2? (LU 16), 1 ? (LU 18); 1 Aug 1964, treehole, R. Martinez and A. Guerra, 3 L (LU 143); 6 Aug 1964, bamboo, R. Martinez and A. Guerra, 14 L(LU 159), 20 L(LU 160), 3 lpd (161-12,13,14), 1 IP (161-13), 3 p (LU 161) [UCLA]. Castries: Bise Quarry, elev. 250 ft, 17 July 1964, treehole, R. Martinez and A. Guerra (LU 46), 3 lp? (46-20,22,23), 2 IP (46-21,24), 3 L, 11 [UCLA]. Cul de Sac Estate, 30 July 1964, chicken bait, R. Martinez and A. Guerra, 2°? (LU 129) [UCLA]. Delgos, elev. 750 ft, 14 July 1964, bamboo, T.H.G. Aitken, R. Martinez and A. Guerra, 3 lpd (10-11,13, 15), 2 Ip? (10-12,14), 1 IP (10-16), 1 P, 7 L(LU 10), 3 lp? (11-13-15), 1 p? (11-101), 3 IP (11-10, 12,18), 1 p, 7 L (LU 11), 3 Ip? (12-10-12), 3 L (LU 12), treehole (LU 13), 2 L, 1 1 [UCLA]. Grande Anse, elev. 175 ft, 20 July 1964, heliconia bract, R. Martinez and A. Guerra, 1 L (LU 62), bamboo (LU 63), 1 Ipdé (63-11), 4 lp? (63-10,12-14), 1 pd (63-101), 2 L, 1 1 [UCLA]. La Croix Junction near Poix Bouche, elev. 500 ft, 20 July 1964, treehole, R. Martinez and A. Guerra (LU 59), 1 Ipd (59-10), 6 L [UCLA]. Marquis Estate, elev. 200 ft, 20 July 1964, fallen cacao pod, R. Martinez and A. Guerra, 1 6, 1 ?, 1 p, 3 L (LU 60), fallen cocoanut (LU 61), 2 lp? (61-10,13), 70 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 1 p? (61-11), 16 L [UCLA]. Piton Flor, elev. 1875 ft, 4 Aug 1964, bamboo, R. Martinez and A. Guerra (LU 146), 1 lpé (146-10), 1 L [UCLA]. Union Agricultural Station, elev. 200 ft, 16 July 1964, bamboo, R. Martinez and A. Guerra (LU 42), 1 Ip? (42-10), 3 IP (42-11,13,14), 10 L [UCLA]. Waterworks Road, 20 July 1964, bromeliads, R. Martinez and A. Guerra (LU 67), 1 p? (67-100) [UCLA]. Micoud: Mahaut, elev. 700 ft, 27 July 1964, heliconia bract, A. Guerra (LU 101), 2 Ipd (101-14,15), 2 Ip? (101-12,13), 1 IP (101-11), 9 L [UCLA] . Mon Repos, elev. 300 ft, 28 July 1964, bromeliads, A. Guerra (LU 104), 1 Ipd (104-11), 6 L [UCLA] . Soufriere: Soufriere, elev. 220-500 ft, 20 July 1964, heliconia bract, R. Martinez and A. Guerra, 2 L (LU 78), bamboo (LU 80), 4 lpé (80-10-13), 2 Ip? (80-14,15), 10 L; 22 July 1964, fallen cacao pod, R. Martinez and A. Guerra (LU 83), 1 lp? (83-11), 3 L [UCLA]. 3 SEXLINEATUS SECTION FEMALES. Medium sized to rather large species, wing 2.7-5.0 mm. Head: Subme- dian and lateral dark patches broad, distinct. Erect scales restricted to occiput. Torus dark brown to almost black, with light scales restricted usually to mesal surface. Pal- pus with or without apical light scales. Thorax: Mesonotum always with distinct lin- ear light pattern, light lines of same color throughout, white, creamy or golden (ex- cept in ecuadoriensis). Acrostichal line not developed but a small patch of light scales present at anterior promontory; inner and outer dorsocentral lines usually long, both reaching scutellum; lateral marginal line usually continuous (broken in Whitmorei Group); median prescutellar line not developed. Paratergite with scales. Acrostichal bristles absent on disc. Pleural scales silvery, white or creamy; ppn and ssp with scales; psp usually with scales (usually absent in Allotecnon Group); mep with separate upper and lower patches. Legs: Midfemur and hindfemur sometimes with distal white spots on anterior surface. All tibiae usually completely dark (ex- cept Sexlineatus and Whitmorei Groups). Tarsal light markings not restricted to seg- ment 1. Wing: Light scales present on remigium in Aurivittatus Group, absent in others. Abdomen: Sternites II-IV with dark scales distally. MALES. Larger claw of foreleg and midleg with submedian tooth only. MALE GENITALIA. Lobes of tergite IX either poorly defined or distinct, bearing a few to many strong to moderate setae. Sidepieces separated from each other at base by a distance less than diameter of aedeagus at that level (greater in Whitmorei Group); tergomesal and/or sternomesal margins with lobes only in Whitmorei Group. Claspette completely absent in Whitmorei Group, represented in other groups by sternal basomesal lobe not connected across midline and bearing 1 apical seta. PUPAE. Cephalothorax: Hair 5-C longer than distance between its base and that of trumpet, strong and always double, at least 2.0 of hair 4-C. Abdomen: Hair 5-IV, V strong, usually double, extending distinctly beyond apex of tergite following; 2- IV-VI usually within basal 0.6 of its tergite; hair 9-III-VI distinctly cephalo-laterad of hair 6; 9-VII moderate to strongly developed, removed considerable distance cepha- lad of hair 6. LARVAE. Stellate hairs always strongly developed. Antenna: Short to moderate; shaft with scattered spicules; hair 1-A usually 2,3-branched. Thorax: Integument with or without spicules. Tubercles of hairs 5-7-P distinctly joined; hair 9-P with more than 4 branches and faintly stellate; 13-P present, distinctly stellate. Abdomen: Integumentary spiculation strong or weak, sometimes confined to dorsal or ventral surface, spicules usually hairlike (spinelike in Allotecnon Group). Hairs 1,2,5,11,13- I, 1,2,5,7,9 (and usually 13)-II-VII stellate; hair 8-VII cephalodorsad or caudodorsad Berlin: Subgenus Howardina Tt of 9-VII. Segment VIII: Hair 1,5-VIII usually stellate. Comb scales in a single row; individual scales long, pointed apically. Siphon: Index 2.0-4.3; integument smooth, imbricate or spiculate. Acus distinct, separate or attached. Pecten teeth with basal denticles. Anal Segment: Saddle with distinct marginal spicules. Hair 1-X moderate, with few branches; 3-X single or double; 2-X multiple. Ventral brush with 5 or 6 pairs of hairs arising from a boss with distinct grid bars. | DISCUSSION. The ornamentation pattern of the adults of the Sexlineatus Sec- tion is superficially so similar in most species that few specimens have been correctly identified in the past and many distinct species were not recognized until in the pres- ent study correlated differences were found in the immature stages. On the whole, the section is a homogeneous one and strongly differentiated from the other 2 sec- tions in all stages as indicated in the keys and diagnosis. As far as is known, members of this section breed only in bromeliad axils. The Sexlineatus Section is primarily associated with the great cordilleras of Cen- tral America and western South America. Apparently only the monotypic Sexlinea- tus Group and the Whitmorei Group penetrate eastward into the lowlands of the Orinoco and the smaller northern South American drainages and no species extends even to the Guianas. There are only small extensions from the Andes down into mod- erate elevations near the headwaters of the Orinoco, Amazon and La Plata basins (fig.1). Only 2 of the 6 groups recognized here for the 19 species are strongly differenti- ated: (1) the Quadrivittatus Group which is confined to Middle America and appears to have retained the most significant primitive features in the section, and (2) the. Whitmorei Group which is known in Colombia, Venezuela and Bolivia but undoubt- edly also occurs in Ecuador and Peru; this is the most derived group in the section. The remaining 4 groups show considerable overlap in many characters although the Sexlineatus and Allotecnon Groups each have a characteristic facies in the adult and larval stages. The Eleanorae Group in many respects appears intermediate between the Quadrivittatus and the Allotecnon Groups. The separation of the Aurivittatus Group from the Eleanorae Group is based only on the presence of light scales on the remigium of the wing of the former; when the immature stages are known the 2 taxo- nomic groups may have to be merged. Two of the groups, Quadrivittatus and Allotecnon, are confined to Middle Ameri- ca where they are sympatric and extend northward to southern and central Mexico. The monotypic Sexlineatus Group overlaps with the above throughout their range in Middle America but is apparently not completely sympatric with them; it further ex- tends along the Pacific coast to the Gulf of Guayaquil and along the southern Carib- bean shores to Trinidad; it is the only species of the section to be found near sea level. The Whitmorei, Eleanorae and Aurivittatus Groups are confined to South America. The dominant Eleanorae Group is primarily Andean and has a wide distri- bution from western Venezuela and Colombia down to Bolivia, with slight intrusions into the upper Orinoco and Amazon basins. The Aurivittatus Group is known only from the upper drainages of the Amazon and La Plata in Bolivia and northern Argen- tina respectively. The Whitmorei Group has been found in Colombia, Venezuela and Bolivia and has small intrusions into the upper Orinoco, Amazon and La Plata drain- ages. Both in Middle and South American groups, some members of a given group seem to occur sympatrically, others are allopatric and still others appear to be ecologically isolated (in the Eleanorae Group) although broadly sympatric. 72 2(1). 3(2). 4(1). 5(4). 6(5). 7(A4). 8(7). Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 KEYS TO GROUPS AND SPECIES ADULTS Midfemur without any indication of a white or silvery spot or streak on an- terior surface above ventral light line (Quadrivittatus Group) . . . . .2 Midfemur with a more or less distinct white or silvery spot or streak on ante- Wor SUriace gbove vet light lime . 8 Pe ee Quadrivittatus Group Outer dorsocentral line not reaching lateral marginal line anteriorly 19. brevis; 20. spinosus Outer dorsocentral line ‘connected to lateral ‘marginal line anteriorly ts Ppn scales predominantly narrow and yellowish; ssp scales in a restricted lin- ear patch not connected to ppn patch; first flagellar segment with dark pedics 9. 1 0, -, 3.027 quadrivittatus Ppn scales predominantly broad and white: ssp scales extending toward ppn patch, the 2 patches usually connected; first flagellar segment with white Oe . 18. lorraineae Antealar area (above paratergite) with numerous dark bronzy scales below narrow supraalar golden line (Sexlineatus and Whitmorei Groups) . . . 5 Antealar area with whitish, yellowish or golden scales only, so that lateral Marginal anc siuptaaiar lines are united .... 8 er a Sexlineatus Group Outer dorsocentral line reaching anterior border of mesonotum; torus with small mesal patch of white or whitish scales. . . . . . .21. sexlineatus Outer dorsocentral line not reaching anterior border of mesonotum, directed outward toward lateral mesonotal line in fossa; torus with a broad ring of white or silvery scales interrupted only dorsally (Whitmorei Group) . .6 Whitmorei Group Hindtarsal white rings on segments 1,2 and 3; anterior end of outer dorsocen- tral line with a patch of broad silvery scales. . . . . . .22. whitmorei Hindtarsal white rings on segments | and 2 only; anterior end of outer dorso- central line without patch of broad silvery scales. . 23. pseudodominicii Base of vein R (remigium) with white or whitish scales . . ..... .8 Base of vein K with dark scales as elsewhere; .. 2 2 ee Aurivittatus Group Hindtarsal white ring on segment 1 only; haltere knob with white scales only . .35. vanemdeni Hindtarsal white eee on ‘sestients 1 2 and - haltere knob with both dark and light seales. 3 5. ; ae Oo ee a Pee ee ke Berlin: Subgenus Howardina 73 9(8). Outer dorsocentral line reaching anterior border of mesonotum; lateral mar- ginal line poorly developed in front of scutal angle, without broadened SCaleS 4 5: Coa SS. QUTIVETalGs Outer dorsocentral line not teachin ariterior ‘border of mesonotum; lateral marginal line strongly developed in front of scutal angle, with broadened SCALES ui aoe Gr we Hinahs ls 5 ee en ne a Eleanorae Group 10(8). Hindtarsus with basal white rings on segments 1 and 3 only. . 27. eleanorae Hindtarsus with basal white rings on segments 1,2, 1-3 or 14... .. 11 11(10).Hindtarsus with basal white rings on segments eS wee Beet. 4 ie Hindtarsus with basal white rings on segments 1,2 or 1-3 only. ie OLS 12(11).Inner dorsocentral line white, usually very narrow; ppn with narrow curved light scales in upper part and broad flat white scales in lower; ssp scale patch extended upward toward ppn broad scales; hindtarsal markings broad, on segment 4 about 0.5 of segment length . . .31. ecuadoriensis Inner dorsocentral line yellowish, usually broadened anteriorly; ppn predom- inantly with moderately broad, curved, light scales throughout; ssp scale patch entirely linear, not extended to ppn scales; hindtarsal markings nar- rower, on segment 4 usually less than 0.2 of segment length aes brevivittatus 13(11).Hindfemur without silvery spot on the anterior surface; ssp scale patch nar- row, linear and not extended toward apn or ppn scale patches. . . . 14 Hindfemur with a very distinct silvery spot on dark portion of anterior sur- face not connected to basal ventral light streak; ssp scale patch short and broad and extended toward apn and ppn scale patches (Allotecnon Group) 16 14(13).Midfemoral light patch or spot at about0.5. .... . . . . 30. osornoi Madfemoral light patch.at about 0.67 of distad 2.6.6. dc nei euetante . > FS 15(14).Outer dorsocentral line strongly broadened anteriorly and joining lateral me- SOUOtal ite 2... ok tis wt ae, merinkellel Outer dorsocentral line not strongly broadened. anteriorly and not joining Valeral TICROMOta) ANG boca cr ee ak a a ae ee Allotecnon Group 16(13).Hindtarsus with basal white rings on segments 1 and 2 only. 20. guatemala Hindtarsus with basal white rings on n seements is. tg ae ee 17(16).Ppn with pure white broad flat scales except for narrow upper border of yel- lowish curved moderately broad scales; psp scales usually present. co 2, Swerrero Ppn predominantly with yellowish, narrow to moderately broad, curved 74 2(1). 3(1). 4(3). 5(4). 6(5). 7(6). 8(7). (5). 10(9). Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 scales throughout except at lower posterior angle where a few broad white flat scales may be present; psp scales absent. . . . . . .24. allotecnon MALE GENITALIA (18. lorraineae,; 25 guerrero; 26 guatemala; 28. marinkellei; and 34. martinezi unknown; 35. vanemdeni not included) Claspette completely absent. . . ta Claspette represented by a small sternomesal lobe bearing a 1 differentiated PO Se ee re ee ne ee Hee hie! Mee Le Tergomesal and sternomesal margins of sidepiece with row of flattened foli- form setae. soo 6.3 . . .22, whitmorei Tergomesal and sternomesal - margins of sidepiece with simple bristles . 23. pseudodominicii Apex of aedeagus in dorsal aspect uniformly sclerotized, smoothly rounded, without indication of apical lateral lobes or apical horns . ae pe § sexlineatus Apex of Aedeaeus deeply lemareiriate and with distinicd long apical horns; or dorsal sclerotization more or less emarginate apically, producing lateral Sletten (ee ee a ee ee Or apa ae Apex of aedeagus with slender horns, bent dorsad . 19. brevis; 20. spinosus Dorsal sclerotization of aedeagus more or less deeply emarginate, producing Riieral Serer Fes er eee Spiniform of clasper short and thick, usually less than 0.25 of clasper length Spiniform of clasper long and slender, usually at least 0.33 of clasper length Aedeagus longer than proctiger ... sayy Catt wr duster iba atte oe Aedeagus shorter than or subequal to proctiger Jenks ie amet staan veers Dis ai etem eT gO Se Inner setae of ninth tergite lobe very markedly flattened, almost leaflike. . . .d1. ecuadoriensis Inner setae of ninth tergite lobe thickened but not markedly flattened . .8 Dorsal sclerotization of aedeagus deeply emarginate, lateral apical lobes con- SPICUOUS “6 os . . 17. quadrivittatus Dorsal sclerotization of aedeagus shallowly emarginate, lateral apical lobes in- er Ra TR nd Oo Oe ane tee aoe Spiniform of clasper about 0.5 of clasper length bate & eleanorae. 33. durivittatus Soiniforn of clasper jess than 0. 4 of clasper POG Make ce Chiat fuer es 2 Dorsal sclerotization of aedeagus deeply emarginate, lateral apical lobes con- RCE a at Cate Wah area ii Mae Me tears 3(2). 4(2). 5(4). 6(5). 7(6). 8(6). 9(8). 2(1). Berlin: Subgenus Howardina 3 Dorsal sclerotization of aedeagus shallowly emarginate, lateral lobes incon- Sei cuss ua ke eae As Os ae er leat i 2 ga ee PUPAE (23. pseudodominicii; 26. guatemala; 31. ecuadoriensis; 32. brevivittatus; 33. aurivittatus; 34. martinezi and 35. vanemdeni unknown; 18. lorraineae and 25. guerrero not included) Trumpet short and broad, index about3.0. . . . . . . .22. whitmorei Trumpet dong and narrow tides over 3.0. ge Oe ee Hairs 10,11,12-C all single . . diag 4 Pi eat i Sig ig a At least one of these hairs branched, usually 10-C. Bae ey ai apt os at ea Ri hae Hair 8-Vi) ondersal surfacts 2 hc ere i ec A ee ie ae eal Hair 8-Vil on ventral surface. Sie 2 ies A ee Ora amelle Hair 8-VII on dorsal surface, strongly developed. . . . . . 27. eleanorae Hay B-V il ow ventral gorivce weak. ae i a Oe ee ee ee Paddle-deeply emarinate-at gir bo. eee eee ee AS Brews Paddle rounded or produced at heir a i2 0) ih Oa A OE er Hair 5-IV-V moderate, not exceeding apex of second tergite following . .7 Hair 5-IV-V very long, exceeding apex of second tergite following . . . .8 Hair 9-VII weak and short, barely reaching lateral angle of tergite . 30. osornoi Hair 9-VIL stronger and longer, extending beyond lateral angle of tergite . 21. sexlineatus Hair 1-II usually double or triple; 9-VIII with 3,4 branches . fa ll ane allotecnon Hair 1 TI usually at least 5-branched: 9-VIII with at least 4 branches . .9 Hair 9-VIII usually at least 8-branched; 9-VII at least 4-branched . ety oie it quadrivittatus Hair 9-VIII usually 6-branched: 9-VII double He eg gue As eo eeEES LARVAE (23. pseudodominicii; 26 guatemala; 31. ecuadoriensis; 32. brevivittatus; 33. aurivittatus; 34. martinezi and 35. vanemdeni unknown) Ventral brush with 6 pairs of hairs ibn resi si ea ae Ventral brush with 5 pairs of hairs . . . TSE ON Quadrivittatus Group Siphon without spicules visible at 10OX; hair 12-C double . . . ... .3 76 32). 4(2). 5(1). 6(5). 7(6). 8(7). 9(6). 10(9). Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Siphon distinctly spiculate at 100X; hair 12-C stellate, with many branches .4 Hair 5-C usually with 5 branches 2... 2 7 geedriviffatus Hait 5-C usually with 3.4 branches .. ........ 4. ...c .. ..d6. lorraineae Hair 5-C usually with more than 10 branches (11-22); ae index about OS sa, “PhD: reves Hair 5-C usually s branched (4- a. siphon index about 2: ie . 20. Spinosus Whitmorei Group Siphon with distinct spicules visible at 1OOX . . . . . . .22. whitmorei Sip) Ort Without spices Vidivic gd) 10UX a ee Hair 15-C usually triple, very short, at most Sait: level of base of mental Diate. al Hair 15-C usually single or r double, moderate i in leneth, reaching level of mid- die Of mental pate (Eicanorde Group). se Sexlineatus Group Thoracic and abdominal integument with dense vestiture of hairlike spicules oo 21, sexlineatus Thoracic integument slibrous at 100X, abdominal with restricted dorsal and ventral patch of small spinelike spicules (Allotecnon Group) .... .8 Allotecnon Group Anal saddle with distinct large caudal marginal spicules ventrad of hair 1-X. . .24. allotecnon Anal saddle without any caudal marginal spicules ventrad of hair 1-X . 2), guerrero Eleanorae Group Siphonal hair 1-S within last pecten tooth; hair 5-C with more than 5 branches <.*~. . . 2/7, eleanorae Siphonal hair 1-S distad of last pecten tooth: hair 5-C single or double .° 10 Peete a ee ie a esome, Tinie oe odors 8 ee A a CO a a 11(10).Siphonal hair 1-S usually single and strongly barbed, rarely double; stellate hairs all shorter than saddle Spee abdominal hairs 1 and 2 usually with about 10-15 branches . . . . 28. marinkellei Siphonal hair 1-S usually triple or - double: many stellate hairs distinctly long- er than saddle, abdominal hairs 1 and 2 usually with 20 or more branches — 29. leei Quadrivittatus Group Berlin: Subgenus Howardina | qi FEMALES. Head: Median light line on vertex with narrow scales. Proboscis dis- tinctly longer than forefemur. Torus with a restricted mesal scale patch. Thorax: Mesonotal light lines usually creamy to golden; inner dorsocentral line complete; outer dorsocentral line either complete or incomplete, not reaching anterior margin; lateral marginal line continuous; scales on anterior promontory golden. Antealar area above paratergite with golden scales. Paratergite with a moderate scale patch extend- ing entire sclerite length. Upper ppn entirely covered with light scales; upper stp patch quadrate. Legs: Distal white spot on anterior surface of midfemur and hindfe- mur absent. Tibiae completely dark. Wing: Veins entirely dark scaled. Abdomen: Basomedian tergal light markings absent. MALES. Palpus entirely dark scaled. MALE GENITALIA. Lobe of tergite IX prominent to indistinct, bearing few (3-5) strong or many (8-10) moderate bristles. Sidepieces separated at base by a distance less than width of aedeagus; without any indication of lobes on tergomesal or sterno- mesal margins. Claspette represented by a conical tubercle. Aedeagus shorter than proctiger; apex moderate to deeply emarginate, producing lateral lobes or horns. PUPAE. Cephalothorax: Trumpet moderate, index about 5.5-6.0, not expanded apically; hair 10-C at least 5-branched; 12-C single, rarely double. Abdomen: Hair 5- IV-V moderate to long, usually exceeding apex of second tergite following (except in brevis); hair 9-VII distinctly stronger than hair 6, exceeding lateral angle of tergite. Paddle: Moderate to long; apex usually produced (emarginate in brevis). LARVAE. Head: Hairs 4-7-C moderately pigmented, long, all at least 5-branched; hair 15-C long, exceeding apex of mental plate. Thorax: Roundish in outline, integu- ment densely spiculate, spicules hairlike, simple or branched. Abdomen: Integument strongly uniformly spiculate. Hair 8-VII slightly cephalodorsad of 9-VII. Siphon: In- tegument imbricate or spiculate; acus distinct, separate. Anal Segment: Hair 3-X sin- gle; ventral brush with 6 pairs of hairs. DISCUSSION. The Quadrivittatus Group of 4 species is distinguished by unique features both in the adults (absence of midfemoral and hindfemoral light spots) and in the larvae (ventral brush with 6 pairs of hairs). Since both these characters are primitive ones in the subgenus, this group probably represents the most ancient lin- eage in the section. ) The group is known only in Central America from southern Mexico to central Panama, usually at elevations above 4,000 feet, and is sympatric with the Allotecnon Group (fig.5). A. quadrivittatus is the only widespread species, occupying the entire known range of the group; lorraineae is restricted to the Sierra Madre del Sur and Sierra Madre de Chiapas in Mexico, where it is sympatric with quadrivittatus, spino- sus and brevis are sympatric with quadrivittatus in the southern part of the range, spinosus in the mountains of Costa Rica and western Panama, and brevis apparently only in western Panama. 17. Aedes (Howardina) quadrivittatus (Coquillett) Figs.5 ,8,36,37 1902.Culex quadrivittatus Coquillett, 1902:292. TYPE: Holotype ¢, Chacula (Huchastenange). Guatemala, elev. 6600 ft, 13 June 1902, Gustav Eisen [USNM, 6560]. Aedes (Howardina) quadrivittatus of Bonne and Bonne-Wepster (1925:369); Carpenter, Galindo and Trapido (1952:163); Horsfall (1955:471); Trapido, Galindo and Carpenter (1955:530,537, 78 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 538); Vargas (1956:22); Trapido and Galindo (1957:122,125 ,130); Galindo (1964:34,56); Bel- kin, Schick and Heinemann (1965:28); Forattini (1965 :401-403); Diaz Najera (1966b:63). Aedes quadrivittatus of Dyar and Knab (1907:11); Theobald (1910:486); Picado (1913:353); Howard, Dyar and Knab (1917:852-853); Dyar (1918:73); Kumm, Komp and Ruiz (1940:400, 417); Arnett (1949:238); Galindo, Carpenter and Trapido (1955:163); Stone and Knight (1956: 224); Blanton and Peyton (1958:31). Aedes (Howardina) quadrivittatus in part of Dyar (1928:229-230); Edwards (1932:155); Lane (1939:100; 1953:40); Stone, Knight and Starcke (1959:174). Culex quadrivittatus of Blanchard (1905:628); Theobald (1905:29; 1910:395). FEMALE. Wing: 3.33 mm. Proboscis: 2.16 mm. Forefemur: 1.8 mm. Abdomen: about 3.06 mm. Head: Light decumbent scales golden on median stripe and short or- bital line; sides and venter creamy. Erect scales on occiput dull golden. Palpus about 0.2 of proboscis, with a few white scales apically and mesally on segment 4. Basal flagellar segment with dark scales. Thorax (fig.8): Mesonotal light scales golden; outer dorsocentral line complete, only slightly expanded at fossa, merging with lateral mar- ginal line at level of humeral bristles; lateral marginal line long, extending from hu- meral bristles to wingbase. Median scutellar lobe with broad dark scales laterally and narrow median golden scales from base to apex; lateral lobe with a few linear golden scales. Paratergite with pale golden linear scales. Apn with narrow anterior and broad- er posterior pale golden scales; ppn with only an upper patch of pale golden linear scales; psp patch pale golden, linear; ssp patch with creamy broad scales; other pleu- ral patches white, scales broad with distribution as in the group. Legs: Forefemur and midfemur predominantly dark anteriorly except for a few basal creamy scales, proximal half of posterior surfaces crearny and distal dark; hindfemur creamy up to 0.67 on both sides, rest dark. Foretarsus with a few basal white scales on segments 1,2; midtarsus with short, often incomplete basal white rings on 1,2; hindtarsus with moderate white rings on 1-3; rest of tarsi dark. Haltere: Stem pale, knob with dark scales basally; fringe whitish. Abdomen: Sternites with broad, V-shaped basal creamy patch and rest dark; with a few apical creamy scales on II-V. MALE. Wing: 3.42 mm. Proboscis: 2.43 mm. Forefemur: 1.98 mm. Essentially similar to female in color. Submedian dark area of head narrower than in female. Median scutellar lobe with larger number of golden scales. MALE GENITALIA (fig.36). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with about 3-5 strong bristles. Sidepiece length 3.0 of medi- an width. Apical seta of claspette on a short basal tubercle, broader than long. Clasper about 0.6 of sidepiece, with 1,2 subapical hairs; spiniform short, about 0.25 of clasper. Aedeagus shorter than proctiger; dorsal sclerotization more or less deeply emarginate, producing distinct lateral apical lobes. PUPA (fig.36). Abdomen: 3.56 mm. Trumpet: 0.51 mm; index about o3Se3 Pad dle: 0.81 mm. Readily separated from brevis by the elongate paddle with produced apex; from spinosus by hair 9-VIII usually at least 8-branched. Chaetotaxy as fig- ured, hairs moderately to darkly pigmented. Cephalothorax: Mesonotum between trumpets moderately pigmented, lateral areas deeper; metanotum strongly pigmen- ted; trumpet strongly contrasting. Hair 1(1, 1-3), 2(1, 1-2), 3(1), 4@, 2-3), 5@), 6(1), 7(2), 8(2, 1-3), 9(1), 10(6, 2-7), 11(1, 1-2), 12(1, 1-3). Abdomen: Tergite I strongly pigmented except for usual membranous areas; segments II-IV strongly pig- mented and imbricate, rest progressively lighter caudad. Segment I: hair 1(8-13 pri- mary branches, each with several secondary branches), 2(1, 1-6), 3(1), 4(2, 2-3), 5(2, 1-3), 6(1, 1-2), 7(1, 1-2), 9(1). Segment II: hair 0(1), 1(5, 2-10), 2(1, 1-3), 3(1), 4G, 2-5), 51, 1-2), 6(1), 7C1, 1-2), 9(1). Segment III: hair 0(1), 1(1, 1-3), 2(1, 1-2), 3(1), Berlin: Subgenus Howardina fie 4(1, 1-2), 5(1, 1-2), 6(1), 7(2, 1-3), 8(2, 1-3), 9(1), 10(1), 11(1), 14(1). Segment IV: hair O(1), 1€1, 1-2), 21, 1-2), 3(2), 4(1, 1-2), 5(2), 6(1), 7(1, 1-3), 8(2, 1-3), 9(1), 10(1), 11¢€1), 14(1). Segment V: hair 0(1), 1(1, 1-2), 2(1), 3(1), 41, 1-2), 5(2, 1-2), 6(1), 7(2, 2-4), 8(2, 1-2), 9(1), 10(1), 11(1), 14(1). Segment VI: hair 0(1), 1(1), 2(1), 31), 41), 51, 1-2), 6(1, 1-3), 701, 1-2), 8(2, 1-2), 9(1), 10(1), 1101), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(1, 1-2), 4(1), 51), 6(1, 1-2), 7(1), 81, 1-2), 9(4, 3-6), 10(1), 11(1), 14(1). Segment VIII: hair 0(1), 4(1), 9(9, 7-11), 14(1, 1-2). Paddle: Moderately to deeply pigmented, midrib darker; apex distinctly produced; external margin serrate; hair 1 slender. Male genital lobe extending to 0.4 of paddle, female genital lobe to 0.31. LARVA (fig.37). Head: 0.86 mm. Siphon: 0.97 mm. Anal Saddle: 0.27 mm. Read- ily separated from spinosus and brevis by the siphon without distinct spicules and hair 12-C double; from lorraineae by hair 5-C usually 5-branched. Chaetotaxy as fig- ured, hairs moderately to darkly pigmented. Head: Width subequal to length; pig- mentation moderate except around mouthparts where it is deeper. Mental plate with a strong median tooth and 11 distinct teeth (11-12) on each side, often a smaller de- tached tooth at base. Hair 0(1), 1(1), 3(1), 4(8, 8-11), 5(6, 5-7), 6(7, 5-10), 79, 8- 15), 8(1), 965, 4-6), 10(2), 11°12, 10-17), 12(2, 2-4), 13(1), 14(12, 11-17), 15@3, 2- 3). Antenna: Length about 0.45 of head; pigmentation moderate. All hairs single ex- cept 1-A which is branched (3, 2-3), and at 0.5 from base. Thorax: Integument with simple or branched hairlike spicules. Stellate hairs with many barbed branches; long hairs barbed and with attenuate apices. Prothorax: hair 0(19, 13-26), 1(5, 5-8), 2(1), 3(9, 8-11), 4(3, 3-4), 5(2, 1-3), 6(1), 7(2), 8(16, 16-20), 9(9, 7-10), 10(1), 112, 1-3), 12(1), 13(24, 20-27), 14(38, 38-48). Mesothorax: hair 1(11, 10-16), 2(1), 3(1), 4(1), 5(1), 6(6, 5-6), 7(1), 8(8, 6-9), 9(9, 7-10), 10(1), 11(1), 121), 13(6, 3-8), 14(8, 7- 12). Metathorax: hair 1(14, 11-18), 2(1), 3(4, 3-7), 4(15, 12-20), 5(15, 13-20), 6(1), 7(8, 7-10), 8(7, 4-8), 9(5, 5-7), 10(1), 11€1), 12(1), 1317, 16-23). Abdomen: Integ- ument and stellate hairs as on thorax. Segment I: hair 1(21, 16-25), 2(21, 19-28), 3(1), 4(1), 5(14, 14-19), 6(3, 3-4), 7(1), 9(4, 3-6), 10(1), 11(18, 16-24), 13(10, 8- 17). Segment II: hair 0(1), 1(18, 17-22), 2(22, 17-28), 3(1), 4(1), 5(18, 16-21), 6(3, 3-5), 7(8, 7-9), 8(1), 9(6, 5-8), 10(1), 11(1), 12(1), 1314, 12-19). Segment III: hair O(1), 1(21, 18-25), 2(18, 16-26), 3(1), 4(1), 5(21, 16-26), 6(1), 7(10, 8-11), 8(1), 9(12, 7-15), 10(1), 11(1), 12(1), 13(20, 17-21), 14(1). Segment IV: hair 0(1), 1(20, 18-27), 2(17, 16-27), 3(1), 4(1), 5(21, 15-24), 6(1), 7(8, 8-10), 8(1), 9(16, 10-16), 10(1), 11(€1), 12€1), 13(20, 15-21), 14(1). Segment V: hair 0(1), 1(21, 19-27), 2(16, 16-22), 3(1), 4(1), 5(20, 14-20), 6(1), 7(9, 8-12), 8(1), 9(16, 11-19), 10(1), 11(1), 12(1), 13(18, 14-20), 14(1). Segment VI: hair 0(1), 1(17, 17-23), 2(18, 14-21), 3(1), 4(1), 5(17, 15-19), 6(1), 7(9, 8-13), 8(1), 9(15, 10-16), 10(1), 11(1), 12(1), 1315, 11-15), 141). Segment VII: hair 0(1), 1(18, 16-21), 2(16, 14-19), 3(1), 4(1), 5(16, 9-19), 6(6, 4-7), 7(2, 2-3), 8(2, 1-3), 9(9, 8-14), 10(1), 11(1), 12(1), 13(4, 3-5), 14(1). Segment VIII: Comb scales 16(14-20), deeply pigmented, elongate; lateral fringes — not extending to apex. Hair 0(1), 1(11, 9-14), 2(1), 3(8, 6-10), 4(1), 5(5, 5-6), 14(1). Siphon: Index about 4.0-4.2; integument imbricate, moderately to deeply pigmen- ted, with darker basal ring. Pecten with 17(13-18) teeth extending to 0.4 and termi- nating before hair 1-S; individual tooth long, pointed, with basal denticles. Hair 1(3, 3-4), 2(1), 3-5 not visible, 6(1), 7(1), 8(2, 1-2), 9(1). Anal Segment: Saddle deeply pigmented and imbricate; caudal margin with spines of varying length, long ones pre- dominating. Gills long, subequal, as long as ventral brush. Hair 1(2), 2(5, 4-5), 3(1), 4a(4, 3-4), 4b(3, 2-3), 4c(3, 2-4), 4d(3, 2-4), 4e(3, 3-4), 4£(3, 2-4). SYSTEMATICS. A. quadrivittatus is the only species occupying the entire range 80 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 of the Quadrivittatus Group from the southern end of the Sierra Madre Oriental in Mexico to central Panama. I have seen no specimen of quadrivittatus from eastern Panama and all material from South America, formerly reported as this species, actu- ally belongs to new species in the Eleanorae Group. A. quadrivittatus is readily dif- ferentiated in all stages from the 3 other members of the group which have restricted distributions, all peripherally sympatric with quadrivittatus. Its closest relative is un- doubtedly lorraineae (see). Very little variation is apparent throughout the wide known distribution of quadrivittatus except at the northern end in Mexico where the light thoracic scaling of the adults tends to be more yellowish or creamy. No sig- nificant population differentiation in the immature stages has been detected in the relatively large sample examined. BIONOMICS. A. quadrivittatus apparently breeds only in epiphytic bromeliads, primarily in cloud forest areas at elevations above 4,000 feet. Females are readily at- tracted to human bait and display a distinct preference for feeding in the forest cano- py (Carpenter, Galindo and Trapido 1952:163). Recently an unidentified arbovirus has been isolated from quadrivittatus in Panama (Galindo 1964:34). DISTRIBUTION (fig.5). Cordilleras of Middle America in Mexico (southern part of Sierra Madre Oriental and eastern end of Sierra Madre del Sur), Guatemala, Hon- duras, El Salvador, probably Nicaragua, Costa Rica and Panama (western part) at elevations varying from about 4000 ft to 8400 ft. Material examined: 321 specimens; 15 6, 42 9, 43 pupae, 221 larvae; 31 individual rearings (6 pupal, 13 larval, 12 incom- plete). COSTA RICA. Alajuela: Poasito, 14 Nov 1962, C.L. Hogue and W.A. Powder (CR 23), 1 Ip? (23-105), 1 L [UCLA]. Volcan Poas, 1 p? (156), 1 ? [USNM] ; Summit, 9 July 1963, C.L. Hogue, 1 L(CR 145) [UCLA]. Cartago: Elev. 1500 m, C. Picado, 2 9? [USNM]. Cervantes, 9 Nov 1962, J.N. Belkin, C.L. Hogue and W.A. Powder, 3 L (CR 13) [UCLA]. Cipreses, 4 6, 29,9 L, 31 [USNM]. La Sierra, elev. 6300-8400 ft, 24 Nov 1962, C.L. Hogue and W.A. Powder, 1 IP (49-107), 1 6,7 L (CR 49), 1 Ipé (51-105), 1 lp? (51-103), 2 IP (51-107,110), 23,6 P, 1 p, 25 L(CR 51), 1 Ipé (52-105), 2 IP (52-106,108), 12 L(CR 52), 3 Ipd (53-101-103), 1 P, 2 L(CR 53), 1 Ipd (54- 103), 1 Ip? (54-106), 5 IP (54-101,102,104,105,107), 3 P, 1 p, 117 L(CR 54) [UCLA]. EL SALVADOR. Los Esesmiles, H W. Kum, 2k [USNM]. GUATEMALA. Chimaltenango: San Andres Semetabaj, 17 July 1965, iCowsill (GUA 50), 1 lp? (50-10) [UCLA]. Solola, 30 June 1964, J.N. Belkin and J.E. Zavortink (GUA 15), 2 Ipd (15-10, 12), 1 Ip? (15-11), 2 1P (15.13.14), 3 L [UCLA] . Huehuetenango: Chacula, elev. 6600 ft, 13 June A a Eisen, type series, 6 2 [USNM] . Locality not specified: Sept 1902, G. Eisen, 1 9; 14,39 HONDURAS. Tegucigalpa, 17 July 1945, 1 lp [USNM]. MEXICO, Chiapas: Paval, 28 A (Diaz Najera 1966b:61). Teopisca, 19 Aug 1964, E. Fisher and D. Verity, 10 L (MEX 132) [UCLA]. Triunfo, 125 A (Diaz Najera 1966b:61). Morelos: (Vargas 1956:22). Oaxaca: Ixtlan de Juarez, Vivero Rancho Teja, elev. 7200 ft, 12 Aug 1966, D.A. Schroe- der (MEX 430), 1 pd (430-105), 4 p? (430-101-104), 12 9 (MEX 433) [UCLA]. Tamaulipas: Ran- cho del Cielo, 4 Apr 1963, Trevino, 2 2 [USNM]. Veracruz: Banderilla, Feb 1945, G. Lissmann, 2g [UCLA]. PANAMA. Bocas del Toro: near Almirante, 1952-53, 2297 ? (Trapido and Galindo 1957:122, 125). La Zorra, elev. 4400 ft, 5 Apr 1963, A. Quinonez, 2 ? (PA 174), 1 9? (PA 175) [UCLA]. Panama: Cerro Campana, 28 Aug 1963, A. Quinonez, 1 9 (PA 537) [UCLA]. 18. Aedes (Howardina) lorraineae Berlin, n.sp. Figs.5 ,8,38 Berlin: Subgenus Howardina 81 TYPES: Holotype ? (MEX 433), Vivero Rancho Teja, Ixtlan de Juarez, Oaxaca, Mexico, 7200 ft, biting collection, 12 Aug 1966, D.A. Schroeder [USNM]. Paratypes: 7 2 (MEX 433), same data as holotype [USNM, BM, UCLA]. This species is dedicated to the memory of Mrs. Lorraine Belkin. FEMALE. Wing: 3.85 mm. Proboscis: 2.62 mm. Forefemur: 2.19 mm. Abdomen: about 3.15 mm. Generally similar to quadrivittatus and other members of the group, differing primarily in the following. Head: Light decumbent scales on median stripe and short orbital line creamy. Erect scales creamy. Palpus about 0.2 of proboscis, with a few white scales on mesal surface of segment 4. Basal flagellar segment with a few white scales on dorsomesal surface. Thorax (fig.8): Mesonotal light lines creamy to pale golden; outer dorsocentral line complete, broadly joined anteriorly in fossa to entire lateral prescutal area and reaching to scutellum; lateral marginal line shorter, extending from scutal angle to wingbase. Narrow light scales on median scutellar lobe creamy; lateral lobes with a few narrow creamy scales. Paratergite and psp with mod- erately broad, white scales; apn patch creamy; upper ppn with predominantly broad creamy scales; lower ppn white, adjoining ssp patch; other pleural patches white, dis- tributed as in the group. Legs: Forefemur and midfemur predominantly dark anteri- orly except for scattered creamy scales up to proximal 0.5, proximal 0.5 of posterior surfaces creamy, extending distally as narrow line, rest dark; hindfemur creamy prox- imally to 0.6 on both sides and rest dark. Foretarsus with basal white scales on seg- ment 1; midtarsus with short incomplete white rings on 1,2; hindtarsus with moder- ately broad white rings on 1-3. Haltere: Stem pale, knob predominantly white, with a few basal dark scales. Abdomen: Sternites with broad V-shaped basal white patches; apical white scales numerous on II-IV, few on V and absent on distal segments. MALE unknown and PUPA imperfectly known. LARVA (fig.38). Head: 0.81 mm. Siphon: 0.87 mm. Anal Saddle: 0.27 mm. Diag- nostic characters as in the key. Readily separated from quadrivittatus by hair 5-C usually only 3,4-branched; from brevis and spinosus by the absence of spicules on siphon. Head: Mental plate with a strong median tooth and 11 distinct teeth (10-12) on each side, often a small detached tooth at base. Hair 0(1), 1(1), 3(1), 4(7, 5-11), 5(3, 3-4), 6(5, 5-6), 7(6, 6-9), 8(1), 9(2, 1-3), 10(2), 1111, 9-17), 12(@, 1-2), 13(1), 14(8, 6-13), 15(2, 2-3). Antenna: Length about 0.33 of head; uniformly moderately pigmented, with a few scattered spicules; hairs simple except 1(2, 2-3). Thorax: In- tegument predominantly with simple hairlike spicules. Stellate hairs with many un- equal barbed branches; long hairs barbed and with attenuate apices. Prothorax: hair O(16, 13-17), 1(6, 4-6), 2(1), 3(8, 6-10), 4(4, 2-4), 5(2), 6(1), 7(2), 816, 14-19), 9(8, 7-9), 10(1), 11@, 1-2), 12(1), 13(@22, 19-27), 14(40, 38-56). Mesothorax: hair 1(11, 10-18), 2(1), 31), 4(€1), 51), 6(6, 4-6), 7(1), 8(7, 6-8), 9(6, 6-9), 10(1), 11d, 1-2), 12(1), 13(6, 5-9), 14(7, 6-9). Metathorax: hair 1(13, 10-18), 2(1), 3(4, 4-6), 4(10, 10-17), 5(13, 8-16), 6(1), 7(7, 4-8), 8(6, 6-9), 9(5, 4-7), 101), 11€1, 1-2), 12(1), 13@20, 14-22). Abdomen: Long and stellate hairs as in thorax. Segment I: hair 1(18, 15-21), 2(16, 14-22), 3(1), 4(1), 5(12, 10-16), 6(3, 2-3), 71), 9(5, 4-7), 101), 11(17, 14-21), 13(15, 9-21). Segment II: hair 0(1), 1(17, 13-20), 2(20, 12-23), 3(1), 4(1, 1-2), 5(16, 13-19), 6(3), 7(8, 5-10), 8(1), 9(6, 6-9), 10(1), 11(€1), 12(1), 13(14, 11-17). Segment III: hair 0(1), 1(20, 15-22), 2(16, 14-21), 3(1), 4(€1), 5(16, 13-20), 6(1, 1-2), 7(10, 8-11), 8(1), 9(10, 8-16), 10€1), 11(1), 121), 13(16, 16-19), 14(1). Segment IV: hair 0(1), 1(20, 15-23), 2(19, 15-19), 3(1), 4(1), 5(16, 14-22), 6(1, 1-2), 7(10, 7-12), 8(1), 9(12, 10-18), 10(1), 11¢€1), 121), 13(17, 13-21), 14(1). Segment V: hair 0(1), 1(19, 14-21), 2(18, 12-18), 3(1), 4(1), 5(16, 11-17), 6(1, 1-2), 7(11, 8- 11), 8(1), 9(12, 10-17), 10€1), 11(1), 12(1), 13(15, 12-19), 14(1). Segment VI: hair 82 | Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 O(1), 1018, 12-23), 2(17, 12-18), 3C1), 4(1), 5(16, 12-17), 6(1, 1-2), 7010, 8-12), 8(1), 9(13, 8-18), 10(1), 11€1), 12(1), 1312, 10-19), 14(1). Segment VII: hair 0(1), 1(18, 13-23), 2(17, 10-18), 3(1), 4(1), 5(16, 13-20), 6(4, 3-5), 7(2, 2-3), 8(2, 2-4), 9(8, 6-11), 10(1), 11(1), 121), 13(4, 3-4), 14C1). Segment VIIT: Comb scales usually 16(14-20), moderately pigmented, their lateral fringes not extending to apex. Hair O(1), 1(8, 7-16), 2(1), 3(8, 7-9), 41), 5(6, 5-8), 14(1, 1-2). Siphon: Index 3.3-3.6; integument imbricate, moderately pigmented, with darker basal ring. Pecten teeth usually 16(13-18), extending to 0.4. Hair 1(4, 3-5), 2(1), 3-5 not visible, 6(1), 7(1), 8(2, 1-2), 9(1). Anal Segment: Saddle moderately pigmented. Gills subequal, narrow, as long as ventral brush. Hair 1(2, 2-3), 2(5, 4-6), 3(1), 4a(3, 3-4), 4b(3, 2-3), 4c(3, 2-3), 4d(3, 2-3), 4e(3, 2-3), 4f(3, 2-3). SYSTEMATICS. A. lorraineae is undoubtedly closely related to quadrivittatus. It is differentiated in the female most conspicuously by the white scales on the first fla- gellar segment and generally whiter scales on the mesonotum and pleura. The associ- ation of the immature stages with the females is presumptive only since all the rear- ings are incomplete. The larva described here, although very similar to quadrivittatus, is distinct from those of Mexican and other populations of the latter in the smaller number of branches in head hair 5. The pupal stage is also in general similar to quad- rivittatus but cannot be described at this time since only a few imperfect whole pu- pae from incomplete rearings are available. A. lorraineae is known at present only from the Sierra Madre del Sur and the Sierra Madre de Chiapas in Mexico at eleva- tions above 7,000 feet. In the eastern part of this range, in the states of Oaxaca and Chiapas, lorraineae is sympatric with quadrivittatus. In one biting-landing collection (MEX 433) in Oaxaca where the 2 species were taken in approximately equal num- bers there is no indication of intergradation between lorraineae and quadrivittatus. This supports my interpretation that lorraineae is a distinct species and not a geo- graphical isolate of quadrivittatus. BIONOMICS. Immature stages of lorraineae have been collected in epiphytic bro- meliads only. Females of this species are apparently readily attracted to humans. DISTRIBUTION (fig.5). Sierra Madre del Sur and Sierra Madre de Chiapas in Mex- ico at elevations above 7000 ft. Material examined: 75 specimens; 11 °, 5 pupae, 59 larvae; 5 incomplete individual rearings. MEXICO. Chiapas: 27 Mar 1961, R.B. Eads, 1 L [USNM]. Guerrero: Omilteme, 8 Sept 1965, D.A. Schroeder, 3 9 (MEX 361A), 3 IP (362-21-23), 24 L (MEX 362) [UCLA]. Oaxaca: Ixtlan de Juarez, El Punto, elev. 7700 ft, 13 Aug 1966, D.A. Schroeder, 4 L (MEX 434); Vivero Rancho Teja, elev. 7200 ft, 12 Aug 1966, D.A. Schroeder, 2 IP (428-10,11), 9 L (MEX 428), 4 L (MEX 429), 9 L (MEX 430), 3 L (MEX 431), 8 9 (MEX 433, type series) [USNM, BM, UCLA]. 19. Aedes (Howardina) brevis Berlin, n.sp. Figs.5 ,9 39,40 TYPES: Holotype ¢d (PA 171-104) with associated pupal skin and genitalia slide, La Zorra, Chiriquisito, Chiriqui Grande, Bocas del Toro, Panama, 4400 ft, 5 Apr 1963, A. Quinonez [USNM]. Allotype ? (PA 171-102) with associated pupal skin, same data as holotype [USNM]. Paratypes: 1 19 (171-105), 2 IP (171-101,103), 2 P, 18 L, same data as holotype [UCLA]. FEMALE. Wing: 3.78 mm. Proboscis: 2.7 mm. Forefemur: 2.34 mm. Abdomen: about 3.1 mm. Generally similar to quadrivittatus and other members of the group; differing primarily in the following. Head: Palpus about 0.15 of proboscis; with dis- Berlin: Subgenus Howardina 83 tinct apical white scales on segment 4. Thorax (fig.9): Outer dorsocentral line incom- plete, arising at anterior 0.33 and reaching to scutellum; lateral marginal line long, extending from anterior promontory to wingbase, slightly curved inwards at lateral prescutal area. Lateral scutellar lobes with a few narrow dark scales. Paratergite and psp with narrow creamy scales; other pleural scale patches as in quadrivittatus, all dull white. Legs: Forefemur dark anteriorly except for a few basal dingy scales, prox- imal 0.5 of posterior surface dingy white, extending distally as narrow line bordered by dark scales; midfemur dark anteriorly, proximal 0.6 of posterior surface dingy white, distal dark; hindfemur dingy white proximally to 0.67 on both sides, rest dark. Foretarsus with basal white scales on segments 1,2; midtarsus with short in- complete white rings on 1,2; hindtarsus with broader white rings on 1-3, last covering about 0.65 of segment. Haltere: Stem pale; knob predominantly white, with a few dorsal dark scales. Abdomen: Sternites with broad V-shaped basal creamy patches; apical light scales apparent on II, III. MALE. Wing: 3.33 mm. Proboscis: 2.89 mm. Forefemur: 2.28 mm. Essentially similar to the female in color. MALE GENITALIA (fig.39). As figured; diagnostic characters as in the key. Ninth tergite lobe indistinct, with 8-10 moderate bristles. Sidepiece length about 3.0 of me- dian width. Clasper about 0.5 of sidepiece, with 1,2 subapical hairs; spiniform short, length about 0.25 of clasper. Apex of aedeagus with slender horns bent dorsad. PUPA (fig.39). Abdomen: 3.78 mm. Trumpet: 0.54 mm; index about 6.0. Paddle: 0.7 mm. Readily separated from other species of group by the deeply emarginate paddle. Chaetotaxy as given below, hairs moderately to darkly pigmented. Cephalo- thorax: Mesonotum lightly pigmented between trumpets, lateral areas darker; meta- notum moderately pigmented. Hair 1(1, 1-2), 2(1), 3C1), 4(2), 5(2), 6(1), 7(2, 1-2), 8(2, 2-3), 9(1), 10(12, 12-14), 111), 12C1, 1-3). Abdomen: Tergite I uniformly mod- erately pigmented except for usual membranous areas; other segments moderate, pig- mentation progressively lighter caudad. Segment I: hair 1(10-20 primary branches, each with secondary branches), 2(2, 1-4), 3(1), 4(3, 2-4), 5(3, 2-5), 6(1), 7(1, 1-2), 9(1); in some hair 10(1) present. Segment II: hair 0(1), 1(16, 12-20), 21, 1-2), 3(1), 4(3, 1-3), 5(2, 1-3), 6(1), 7(1), 9(1), 11(1). Segment III: hair 0(1), 1(2, 2-3), 2(1), 3(1), 41, 1-2), 5(1, 1-5), 6(1), 7€1, 1-2), 8(1), 91), 10(1), 11¢1), 14(1). Segment IV: hair 0(1), 1(2), 2(1), 3(2, 1-2), 4(1), 5(2), 6(1), 701), 81), 91), 10(1), 1101), 14(1). Segment V: hair 0(1), 1(2, 1-2), 2(1), 3(1), 4(1), 5(2), 6(1), 7@, 1-3), 8(1), 9(1), 10(1), 11(1), 14(1). Segment VI: hair 0(1), 1(1, 1-2), 21), 301), 4(1), 5(), 6(1), 7(2, 1-6), 8(1), 9(1), 10€1), 1101), 14(1). Segment VII: hair 0(1), 1(1, 1-2), 2(1), 3(1), 4(1), 5(1), 6(2, 1-3), 7(1), 8C1, 1-2), 9(7, 5-8), 10(1), 11(1), 141). Seg- ment VIII: hair 0(1), 4(1), 9(11, 11-14), 14(1). Paddle: Uniformly lightly pigmented, midrib slightly darker; deeply emarginate; external margin serrate; hair 1-P single. Male genital lobe extending to about 0.5 of saddle, female genital lobe to 0.35. LARVA (fig.40). Head: 0.96 mm. Siphon: 0.6 mm. Anal Saddle: 0.3 mm. Readi- ly separated from other members of the group by hair 5-C usually with 18 branches (11-22) and short siphon with an index of about 2.0. Head: Mental plate with a strong median tooth and 10 distinct teeth (9-10) on each side, often a small detached tooth at base. Hair 0(1), 1(1), 3(1), 4(16, 13-20), 5(18, 11-22), 6(12, 9-19), 7(15, ~ 11-21), 8(4, 1-7), 9°09, 6-18), 10(6, 3-8), 11(24, 18-25), 12(8, 5-12), 13(1), 14Q19, 15-22). 15(2, 2-4). Antenna: Length about 0.35 of head; shaft uniformly moderate- ly pigmented, with a few scattered spicules; hairs simple except 1(2, 2-3). Thorax: Integument with simple and branched, hairlike spicules. Stellate hairs with many un- equal barbed branches; long hairs barbed and with attenuate apices. Prothorax: hair 84 : Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 0(24, 19-24), 1(12, 10-14), 2(1, 1-2), 3(14, 13-23), 4(4, 3-6), 5@, 2-3), 6(1), 7(3, 2- 3), 8(21, 16-26), 9(7, 6-12), 10(1), 11(@2, 1-2), 121), 13(30, 24-31), 14(38, 34-43). Mesothorax: hair 1(21, 15-33), 2(1), 31), 4(1), 5(1, 1-2), 6(6, 6-9), 7(1), 8(7, 5-7), 99, 7-10), 10(1), 11€1, 1-2), 12(1), 13(16, 9-16), 14(16, 15-21). Metathorax: hair 1(28, 22-30), 2(1), 3(16, 12-19), 4(5, 4-5), 5(12, 11-15), 6(1), 7(6, 4-9), 8(14, 10- 18), 9(5, 5-8), 10€1), 111), 12(1), 1319, 19-35). Abdomen: Stellate hairs as in tho- rax. Segment I: hair 1(29, 29-32), 2(22, 18-31), 3(1), 41, 1-3), 5(18, 14-23), 6(3, 1- 4), 7(1), 9(7, 5-8), 10(1), 11022, 17-22), 13(26, 23-27). Segment II: hair 0(1), 1(26, 24-40), 2(22, 15-26), 3(1), 4€1), 5(18, 16-23), 6(3, 3-4), 7(13, 5-14), 8(1), 9(13, 9- 1S), 101), 1101), 121), 13(25, 21-26). Segment III: hair 0(1), 1(26, 24-35), 2(18, 15-22), 31), 4(1), 5(18, 17-22), 6(3, 2-3), 7014, 11-15), 8(1), 9(16, 12-16), 10(1), 11q1), 121), 13(21, 19-25), 14(1). Segment IV: hair 0(1), 1(26, 24-29), 2(16, 16- 20), 31), 41), 519, 15-21), 6(2), 711, 11-18), 8(1), 9(15, 15-18), 10(1), 1101), 12(1), 13(21, 20-34), 14(1). Segment V: hair 0(1), 1(25, 24-37), 2(18, 15-20), 3(1), 4(1), 5(18, 16-21), 6(2, 2-3), 7(12, 9-18), 8(1), 9(16, 14-24), 10(1), 11(1), 12(1), 13(19, 18-34), 14(1). Segment VI: hair 0(1), 1(27, 24-38), 2(18, 15-19), 3(1), 4(1), 5(16, 14-20), 6(2), 7(12, 11-14), 8(1), 9(16, 14-20), 10(1), 111), 1201), 13(20, 17- 22), 14(1). Segment VII: hair 0(1), 1(25, 23-30), 2(17, 14-22), 3(1), 4(1), 5(16, 14- 18), 6(7, 5-7), 72, 1-2), 8(3, 2-4), 9(19, 14-20), 10(1), 11(1), 12(1), 1319, 17-23), 14(1, 1-2). Segment VIIT: Comb scales usually 16(14-17), moderately pigmented, their lateral fringes not extending to apex. Hair 0(1), 1(18, 15-21), 2(1), 3(7, 6-9), 4(1), 5(16, 15-20), 14(1). Siphon: Index about 2.0; integument moderately pigmen- ted, with darker basal ring and with rows of small hairlike spicules. Pecten teeth usu- ally 9(7-16), extending to 0.3. Hair 1(5, 4-6), 2(1), 3-5 not visible, 6(1), 7(1), 8(1, I- 2), 9(1). Anal Segment: Saddle moderately pigmented, with a darker basal ring and with small spicules; caudal margin with spines of varying length. Gills about 2.0 of saddle length, narrow and subequal. Hair 1(2, 2-3), 2(6, 4-6), 3(1), 4a(4, 3-5), 4b(3, 2-3), 4c(3, 3-4), 4d(4), 4e(4, 3-5), 4£(4, 3-5). SYSTEMATICS. The adults of brevis cannot be differentiated from the apparent- ly closely related spinosus but the immature stages of the 2 species are strikingly dif- ferentiated as indicated in the diagnosis and in the keys. The association of all the stages is undoubtedly correct although no complete larval rearings were made. At present, brevis is known only from northern Panama where it has been found in the same locality as quadrivittatus and spinosus but not in the same collections. BIONOMICS. Immature stages of brevis have been collected in epiphytic bromeli- ads at elevations above 4,000 feet. All 5 known adults were reared but it is possible that the 2 females taken in biting-landing collections in Panama provisionally identi- fied as spinosus are actually brevis. DISTRIBUTION (fig.5). Chiriqui mountains of Panama at elevations above 4000 ft. Material examined: 58 specimens; 3 6, 2 2, 10 pupae, 43 larvae; 8 individual rear- ings (1 larval, 4 pupal, 3 incomplete). PANAMA. Bocas del Toro: La Zorra, elev. 4400 ft, 5 Apr 1963, A. Quinonez (PA 171, type se- ries), 1 1d (171-105), 1 pd (171-104), 1 p? (171-102), 2 IP (171-101,103), 2 P, 18 L;6 Apr 1963, A. Quinonez, 18 L (PA 178); 9 Apr 1963, A. Quinonez (PA 183), 1 pd (183-101), 1 p? (183-103), 1 IP (183-102), 3 L [USNM, UCLA]. 20. Aedes (Howardina) spinosus Berlin, n.sp. Figs.5 ,9,41,42 Berlin: Subgenus Howardina 85 TYPES: Holotype ? (CR 13-111) with associated pupal and larval skins, near Cervantes, Car- tago, Costa Rica, 9 Nov 1962, J.N. Belkin, C.L. Hogue and W.A. Powder [USNM]. Paratypes: 4 larvae (CR 13), same data as holotype [USNM, UCLA]. FEMALE. Wing: 3.33 mm. Proboscis: 2.32 mm. Forefemur: 1.8 mm. Abdomen: 3.06 mm. Holotype lacking part of hindtarsus, entire left leg and abdomen. Essential- ly similar to brevis in all details, from which it cannot be separated. The only appar- ent difference is in the pleural scales which are white in spinosus and dingy white in brevis (fig.9). MALE. Wing: 3.5 mm. Proboscis: 2.8 mm. Forefemur: 2.15 mm. Essentially sim- ilar to female in color. MALE GENITALIA (fig.41). Diagnostic characters as in the key. Essentially simi- lar to brevis in all details. PUPA (fig.41). Abdomen: 3.25 mm. Trumpet: 0.5 mm; index about 6.0. Paddle: 0.77 mm. Readily separated from brevis by the rounded or slightly produced paddle; from quadrivittatus by 2-branched hair 9-VII and 5-branched 9-VIII. Hairs moder- ately to darkly pigmented. Cephalothorax: Mesonotum lightly pigmented between trumpets, lateral areas and metanotum slightly darker; trumpets moderately long, strongly contrasting. Hair 10-C with 5 branches; 12-C double. Abdomen: Tergite I moderately pigmented except for usual membranous areas; pigmentation of rest of segments moderate, progressively lighter caudad. Hair 1-II 8-branched; hair 5-IV,V long, double, exceeding apex of second tergite following; hair 9-VII distinctly larger than 6-VII, double, long, exceeding apex of tergite; hair 9-VIII long, 5-branched. Paddle: Uniformly lightly pigmented, midrib slightly darker; apex rounded or slight- ly produced; hair 1-P single; external margin serrate. Female genital lobe extending to 0.38 of paddle. LARVA (fig.42). Head: 0.88 mm. Siphon: 0.88 mm. Anal Saddle: 0.33 mm. Read- ily separated from quadrivittatus and lorraineae by the pilose siphon; from brevis by the longer siphon and hair 5-C usually 5-branched (4-7). Head: Mental plate with a strong median tooth and 11 distinct teeth (11-12) on each side, often a small de- tached tooth at base. Hair 0(1), 1(1), 3(1), 409, 8-11), 5(5, 4-7), 6(8, 6-9), 7(8, 8-11), 8(2, 1-3), 9(7, 5-9), 10(4, 3-5), 11(21, 17-24), 12(8, 4-10), 13(1), 14(16, 15-24), 15(3). Antenna: Length about 0.33 of head; shaft moderately pigmented and with a few scattered spicules; hairs simple except 1(2, 2-3) which is at about 0.6 from base. Thorax: Integument predominantly with branched, hairlike spicules. Stellate hairs with many unequal barbed branches; long hairs with attenuate apices. Prothorax: hair 0(18, 16-20), 1(5, 5-8), 2(1), 3(11, 10-14), 4(3, 3-5), 5(2), 6(1), 7(2), 8(16, 16- 20), 9(6, 4-7), 10(1), 11(2, 2-3), 12(1), 13(31, 20-38), 14(44, 44-60). Mesothorax: hair 1(18, 9-20), 2(1), 3(1), 4(1), 5(1), 6(6, 4-6), 7(1), 8(7, 6-7), 9(7, 4-7), 10(1), 11(2), 121), 1365, 4-7), 14(8, 5-14). Metathorax: hair 1(25, 14-27), 2(1), 3(4, 3-4), 4(18, 14-20), 5(18, 15-20), 6(1), 7(7, 5-8), 8(10, 5-10), 9(5, 4-6), 10(1), 11(1, 1-2), 12(1), 13(20, 18-24). Abdomen: Segment I: hair 1(28, 24-33), 2(25, 22-27), 3(1), 4(1), 5(20, 16-23), 6(3), 7(1, 1-2), 9(6, 6-8), 10(1), 11(22, 19-25), 13(24, 20-28). Segment II: hair 0(1), 1(30, 26-34), 2(24, 21-28), 3(1), 4(1), 5(28, 22-31), 6(3), 7(14, 10-14), 8(1), 9(14, 11-16), 10(1), 1101), 121), 13(29, 26-31). Segment III: hair O(1), 1(30, 22-35), 2(25, 22-29), 3(1), 4(1), 5(28, 22-31), 6(2), 7(16, 10-16), 8(1), 9(20, 11-22), 10(1), 11(1), 12(1), 13(28, 18-32), 14(1). Segment IV: hair 0(1), 1(30, 21-36), 2(27, 20-28), 3(1), 4(1), 5(26, 19-31), 6(2), 7(16, 9-17), 8(1), 9(20, I-22), LOT), 111), 12019, 13@7,; 20-32), 14€1). Segment V; hair 001), 1134, 23- 34), 2(24, 16-28), 3(1), 4(1), 5(26, 20-31), 6(2), 7(16, 9-19), 8(1), 9(21, 16-24), 86 : Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 10(1), 11(1), 12(1), 13(27, 18-28), 14(1). Segment VI: hair 0(1), 1(32, 21-33), 223, 17-27), 3(1), 4(1), 5(27, 19-29), 6(2), 7(17, 11-20), 8(1), 9(20, 15-24), 10(1), 11(), 12(1), 13(20, 16-24), 14(1). Segment VII: hair 0(1), 1(30, 22-34), 2(25, 17-26), 3(1), 4(1), 5€18, 17-24), 6, 6-11), 7(@, 2-3), 8C1), 9@2, 10-23), 10G); 11(1), 120), 13(17, 13-20), 14(1). Segment VIII: Comb scales usually 16(14-18), moderately pig- mented; individual scale long, pointed apically and fringed nearly to tip. Hair O(1), 1(22, 14-26), 2(1), 3(8, 7-9), 4(1), 5(12, 8-16), 14(1). Siphon: Index about 3.2-4.0; moderately pigmented, except for dark basal ring; uniformly covered with rows of small hairlike spicules. Pecten teeth usually 10(10-15), extending to about 0.4 before hair 1-S; individual tooth long, pointed, with basal denticles. Hair 1(5, 4-5), 2(1), 3-5 not visible, 6(1), 7(1), 8(2), 9(1). Anal Segment: Saddle moderately pigmented ex- cept for darker basal ring; covered with small spicules; caudal margin with serrated spines of varying length. Gills long, subequal, as long as ventral brush. Hair 1(2), 2(6, 6-7), 3(1), 4a(5, 4-5), 4b(3, 3-4), 4c(3, 3-4), 4d(4, 3-4), 4e(3, 3-4), 4f(3, 1-3). SYSTEMATICS. A. spinosus is recognized as a distinct species entirely on the basis of striking and constant features of the immature stages as indicated in the diag- nosis. The single individually reared adult (?) shows no obvious differences from the 5 individually reared specimens of brevis. Therefore, the identity of the male and the 5 females listed below is questionable; they could possibly be brevis. A. spinosus is known at present only from areas of cloud forest above 4,000 feet in Costa Rica and Panama. It is sympatric throughout this range with the dominant quadrivittatus. Al- though the 2 species are reported as being associated in 2 collections of immature stages in Costa Rica (CR 13,54), it is not known if they were actually found together in the same bromeliads. In Panama, spinosus has been collected in the same locality as brevis but in this instance the immature stages were not associated in a given col- lection, suggesting an ecological barrier between the 2 species. BIONOMICS. Immature stages of spinosus occur in epiphytic bromeliads. Two fe- males from Panama provisionally identified as spinosus were taken in biting-landing collections. DISTRIBUTION (fig.5). Costa Rica and Panama (Chiriqui mountains) at eleva- tions above 4000 ft. Material examined: 85 specimens; | 6, 6 2, 3 pupae, 75 larvae; 2 individual rearings (1 larval, 1 incomplete). COSTA RICA. Alajuela: Volcan Poas, 14 Apr 1938, 2 1 [USNM]. Cartago: Cervantes, 9 Nov 1962, J.N. Belkin, C.L. Hogue and W.A. Powder (CR 13, type series), 1 Ip? (13-111), 4 L [USNM, UCLA]. Cipreses, 1 9 [USNM]. La Sierra, elev. 6300 ft, 24 Nov 1962, C.L. Hogue and W.A. Pow- der, 55 L(CR 54) [UCLA]. Locality not specified: C. Picado, 1 6,2 9 [UCLA, USNM]. PANAMA. Bocas del Toro: La Sierra, elev. 5400 ft, 8 Apr 1963, A. Quinonez, 5 L (PA 180), 1 9 (PA 181) [UCLA]. La Zorra, elev. 4400 ft, 5 Apr 1963, A. Quinonez, 1 L (PA 172); 6 Apr 1963, A. Quinonez, 1 IP (176-103), 1 P, 1 L (PA 176); 7 Apr 1963, A. Quinonez,5 L(PA 179); _ . 9 Apr 1963, A. Quinonez, 1 9 (PA.184) [UCLA]. Sexlineatus Group FEMALE. Head: Median light line on vertex with narrow scales. Proboscis dis- tinctly longer than forefemur. Torus with restricted mesal scale patch. Thorax: Meso- notal light lines mostly white, rarely creamy; inner and outer dorsocentral lines com- plete, subparallel; lateral marginal line continuous; scales on anterior promontory white. Antealar area above paratergite with dark scales. Paratergite with a moderate scale patch extending entire sclerite length. Upper ppn entirely covered with scales, Berlin: Subgenus Howardina 87 both dark and light scales present; upper stp patch quadrate. Legs: Distal white spot on anterior surface of midfemur and hindfemur present. Tibiae predominantly dark, with light scales in narrow longitudinal lines. Wing: Veins entirely dark scaled. Abdo- men: Basomedian tergal light markings present on III-VII. MALE. Palpus with basomedian light scales on segments 4,5. MALE GENITALIA. Lobe of tergite IX prominent, with 5,6 strong bristles. Side- pieces separated at base by a distance less than width of aedeagus; without any indi- cation of lobes on tergomesal or sternomesal margins. Claspette represented by a conical tubercle. Aedeagus shorter than proctiger; apex smoothly rounded, without any indication of lateral lobes. PUPA. Cephalothorax: Trumpet moderate, index about 5.0, distinctly flared api- cally; hairs 10,12-C usually with a few branches. Abdomen: Hair 5-IV-V moderate, not exceeding the apex of second tergite following; hair 9-VII moderate to strong, distinctly larger than hair 6, exceeding lateral angle of tergite. Paddle: Short; apex rounded. , LARVAE. Head: Hair 4-C weak, lightly pigmented, at least 4-branched; 5-C sin- gle, moderate, strongly pigmented; 6-C heavy, darkly pigmented, triple or double; 7- C moderate, double or triple; 15-C short, barely exceeding base of mental plate. Tho- rax: Roundish in outline; integument strongly spiculate, spicules hairlike, simple. Abdomen: Integument densely spiculate. Hair 8-VII slightly cephalodorsad of 9-VII. Siphon: Integument imbricate; acus distinct, attached. Anal Segment: Hair 3-X dou- ble; ventral brush with 5 pairs of hairs. DISCUSSION. Only 1 species is recognized at this time in the Sexlineatus Group although it is evident, as indicated under the species, that there is considerable dif- ferentiation in some populations. The Sexlineatus Group shows similarities with the Whitmorei Group in adult orna- mentation (dark-scaled antealar area) and with the Allotecnon Group in the larva (head hair 15-C short). It is the most widespread of all the groups in the section and occurs at lower elevations than any other, particularly in the southern part of its range in Trinidad and in lowland Ecuador where it seems to be the only species of Howardina present (fig.4). In the central (Honduras, Panama) and northern (Mexico) portions of its range it occurs at moderate to high elevations and may be sympatric with the Allotecnon and Quadrivittatus Groups. 21. Aedes (Howardina) sexlineatus (Theobald) Figs.4,9 43,44 1901.Stegomyia sexlineata Theobald, 1901:308. TYPE: Holotype ? (102), Agua Santa (St. George), Trinidad, F.W. Urich [BM]. 1907.Stegomyia dominicii Rangel and Romero-Sierra, 1907:48. TYPE: ?, Hacienda El Paraiso on Rio Anauco [Caracas], Venezuela [LU]. NEW SYNONYMY. Aedes (Howardina) sexlineatus of Dyar (1928:229); Edwards (1932:155); Martini (1935:49); Komp (1936a:57); Lane (1939:101); Martinez and Prosen (1950:38-39); Lane (1953:715-716); Hors- fall (1955:471); Martinez and Prosen (1955:23-25); Reyes (1955:74,89); Trapido, Galindo and Carpenter (1955:530,534,537); Vargas (1956:22); Galindo and Trapido (1957:122,124,125, 134); Trapido and Galindo (1957:134); Belkin, Schick and Heinemann (1965:69); Forattini (1965:113,232); Cova Garcia, Sutil and Rausseo (1966a:58,287; 1966b:74,141 334-335). Aedes (Howardina) sexlineata of Bonne and Bonne-Wepster (1925:367-368). 88 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Aedes sexlineatus of Urich (1913:527,528); Howard, Dyar and Knab (1917:847-849); Dyar (1918: 73); Cerqueira (1943b:33); Boshell-Manrique and Osorno-Mesa (1944:173,174,177,178); Del Ponte, Castro and Garcia (1951:240); Aitken (1960:4). Aedes (Soperia} sexlineata of Anduze (1941:14). Stegomyia sexlineata of Giles (1902:377); Theobald (1903b:238,239); Blanchard (1905:260). Gymnometopa sexlineata of Coquillett (1905:183; 1906b:25); Theobald (1907:210; 1910:219); Surcouf and Gonzalez-Rincones (1911:212-213). Haemagogus sexlineata of Dyar and Knab (1906a:166). Aedes (Howardina) dominicii of Bonne and Bonne-Wepster (1925:376); Dyar (1928:231); Ed- wards (1932:155). Aedes (Soperia) dominicii of Belkin, Schick and Heinemann (1965:74); Cova Garcia, Sutil and Rausseo (1966b:80). Stegomyia dominicii of Surcouf and Gonzalez-Rincones (1911:131,134-135; 1912:286-287). FEMALE. Wing: 2.8 mm. Proboscis: 2.2 mm. Forefemur: 1.9 mm. Abdomen: about 2.9 mm. Head: Light decumbent scales white. Erect scales whitish on occiput, dark brown on sides and disc. Palpus about 0.2 of proboscis, white scales on mesal surface and apex of segment 4. Torus light brown laterally, darker mesally and with white scales. Thorax (fig.9): Mesonotal light lines white to dingy white; inner and outer dorsocentral lines complete from anterior margin to scutellum, scales at level of prescutellar area lighter; lateral marginal line short, narrow from scutal angle to wingbase, scales longer, broader at wingbase. Antealar area above paratergite com- pletely dark scaled. Median scutellar lobe with moderately broad dark scales laterally and with median narrow white scales from base to apex; lateral lobes with narrow dark scales. Paratergite scales broad, white. Apn scales white, predominantly broad, a few anterior scales narrower; upper ppn with narrow black scales on upper half, light to creamy narrow and broad scales on lower half, extent of black scales varied in different populations; psp scales distinct, narrow and creamy in front, broad and white behind; other scale patches pure white, distributed as in group. Legs: Forefe- mur dark anteriorly except for basal creamy scales, dark posteriorly except for a nar- row creamy longitudinal line; midfemur mostly dark anteriorly except for basal creamy scales, a narrow mesal white line to middle and a white spot at distal 0.67, proximal 0.6 of posterior surface white, rest dark scaled; hindfemur predominantly dark anteriorly except for broad white line to 0.75 terminating in a white spot, pos- terior surface white to 0.75, rest dark scaled. Foretibia and midtibia predominantly dark except for anterior, narrow, faint white line; hindtibia dark. Foretarsus with white scales on segments 1,2; midtarsus with short incomplete white rings on 1,2, often a few white scales on 3; hindtarsus with broader white rings on 1-3, occasion- ally some white scales on segment 4 and sometimes on 5. Haltere: Stem pale, knob dark basally, fringe scales white. Abdomen: Basomedian tergal scales creamy, silvery on VII. Sternites with broad basal V-shaped pure white patches; apical white scales apparent up to segment V. MALE. Wing: 2.61 mm. Proboscis: 2.43 mm. Forefemur: 1.8 mm. eRe sim- ilar to female in color. MALE GENITALIA (fig.43). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with 5,6 moderate setae. Sidepiece length about 3.0 of medi- an width. Clasper about 0.5 of sidepiece, spiculate in basal 0.33. Aedeagus short, apex uniformly sclerotized, smoothly rounded, without indication of apical lateral lobes. PUPA (fig.43). Abdomen: 3.6 mm. Trumpet: 0.43 mm; index about 5.0. Paddle: 0.6 mm. Diagnostic characters as in the key. Chaetotaxy as given below, hairs moder- Berlin: Subgenus Howardina 89 ately to darkly pigmented. Cephalothorax: Mesonotum moderately pigmented be- tween trumpets, lateral areas and metanotum slightly darker; trumpets contrasting. Hair 1(1), 2(1), 3(1), 4(3, 2-5), 5(2, 2-3), 6(1, 1-2), 7(2, 2-3), 8(2, 1-4), 9(1), 10(4, 1-7), 111, 1-2), 12(3, 1-4). Abdomen: Tergite I uniformly strongly pigmented ex- cept for usual light membranous area; segments II-VIII with imbricate integument, strongly pigmented, progressively lighter caudad. Segment I: hair 1(6-13 primary branches, each with many secondary ones), 2(1, 1-2), 3(1, 1-2), 4(2, 2-4), 5(3, 1-6), 6(1, 1-2), 71, 1-4), 9(1, 1-3), 10(1-2; present in some), 11(1-4, present in some). Segment IT: hair 0(1), 1(2, 1-7), 2(1, 1-2), 31, 1-2), 4(2, 1-4), 5(1, 1-2), 6C1, 1-2), 7(1, 1-2) present in some, 9(1), 10(1, present in some), 11(1-2, present in some). Seg- ment III: hair 0(1), 1(3, 1-5), 21, 1-3), 3d, 1-2), 4(2, 1-3), 5(1, 1-2), 6(1), 7(2, 1-3), 8(3, 2-4), 9(1), 101, 1-3), 11(1), 14(1). Segment IV: hair 0(1), 1(2, 2-5), 2(1), 3(2, 2-4), 4(2, 1-2), 5(2), 6(1), 7(2, 1-2), 8(2, 1-3), 9(1), 10(1), 11(1), 14(1). Segment V: hair 0(1),°1¢2,.1-3), 2@h, 1-2);:301; 1-3)40; 1-3); 502), 601), 102 1-4); 8, 13), 9(1), 10(1), 11(1), 14(1). Segment VI: hair 0(1), 1(1, 1-2), 2(1), 3(1), 41, 1-2), 5, 1-2), 6(1), 7C1, 1-2), 8(2, 1-2), 9(1), 101, 1-2), 11(1), 14(1). Segment VII: hair 0(1), I(1, 1-2), 2(1), 31), 4€1), 51, 1-3), 6(2, 1-3), 7(1), 8(3, 1-4), 9G, 2-5), 10(1), 11(1), 14(1). Segment VIII: hair 0(1), 4(1), 9(6, 4-9), 14(1). Paddle: Uniformly moderately pigmented, midrib darker; external margin serrated, apex smooth; hair 1-X slender. Male genital lobe extending to 0.59 of paddle, female genital lobe to 0.41. LARVA (fig.44). Head: 0.76 mm. Siphon: 0.65 mm. Anal Saddle: 0.27 mm. Chae- totaxy as given below, hairs moderately to darkly pigmented. Head: Mental plate with a strong median tooth and 9-11 distinct teeth on either side, often a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 4(7, 4-8), 5(1, strong), 6(3, 2-3), 7(3, 2-3), 8(1), 9(2, 2-3), 10(2, 1-2), 11(17, 10-19, stellate), 12(1), 13(1), 14(12, 12-24, stellate), 15(2, 2-4). Antenna: Length about 0.3 of head; pigmentation moderate; shaft with a few scattered denticles. All hairs except 1-A single; hair 1-A (2, 2-3) about 0.6 from base. Thorax: Integument strongly pilose, spicules simple, hairlike; apices of long hairs attenuate and weakly barbed. Stellate hairs with many barbed branches. Pro- thorax: hair 0(23, 19-25), 1(5, 2-8), 2(1, 1-2), 3(20, 18-26), 4(1, 1-3), 5(1), 6(1), 7(2, 2-3), 8(20, 15-23), 9(17, 16-24), 10(1), 11(1, 1-2), 12(1), 13(20, 18-29), 14(32, 30- 42). Mesothorax: hair 1(11, 8-15), 2(1), 3(1), 4(1), 5(1), 6(4, 3-5), 7(1), 8(5, 4-6), 9(5, 4-6), 10(1), 11(2, 1-4), 121, 1-2), 13(17, 12-21), 14(16, 10-20). Metathorax: hair 1(21, 9-21), 2(1), 3(8, 3-9), 4(13, 12-18), 5(11, 9-16), 6(1), 7(4, 3-5), 8(17, 12- 19), 9(3, 2-5), 10(1), 11(2, 2-4). 12(1), 13(22, 16-26). Abdomen: Integument and stellate hairs as on thorax. Spicules as on thorax, but longer and more profuse on ventral side of segments. Segment I: hair 1(20, 12-21), 2(20, 18-25), 3(1), 4(1), 5(15, 13-20), 6(2, 2-3), 7(1), 9(7, 5-10), 10(1), 11(20, 12-23), 13(15, 9-16). Segment II: hair O(1), 1(20, 16-26), 2(20, 16-24), 3(1), 4(1, 1-2), 5(16, 12-17), 6(2, 2-3), 7(9, 5- 11), 8(1), 9(10, 6-13), 10(1, 1-2), 11€1, 1-2), 12(1), 13(14, 12-18). Segment III: hair O(1), 1(20, 16-23), 2(20, 9-24), 3(1), 4(1), 5(14, 14-20), 6(2), 7(14, 11-17), 8(1), 9(14, 10-18), 10(1), 11€1), 12(1), 13(13, 11-17), 14(1). Segment IV: hair 0(1), 1(20, 12-21), 2(22, 13-25), 3(1), 4(1), 5(15, 12-20), 6(2), 7(13, 10-15), 8(1), 9(18, 11-19), 101), 1101), 121), 13C11, 9-17), 141). Segment V: hair 0(1), 1(17, 12-21), 2018. 17-24), 3(1), 4(1), 5(15, 13-20), 6(2), 7(12, 10-15), 8(1), 9(16, 13-19), 10(1); 11(1), 12(1), 13(10, 9-14), 14(1). Segment VI: hair 0(1), 1(17, 12-20), 2(18, 17-22), 3(1), 4(1), 5€16, 13-17), 6(1, 1-2), 7(11, 8-12), 8C1), 9(17, 11-19), 10(2), LTC y, 12). 13(12, 8-15), 14(1). Segment VII: hair 0(1), 1(18, 12-18), 2(18, 15-21), 3(1), 4(1), 5(13, 11-17), 6(8, 8-12), 7(1), 8, 8-10), 9(12, 12-21), 10(1), 11(1), 121), 13(9, 7- 12), 14(1). Segment VII: Comb scales in a single row, usually 10(8-14) in number; 90 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 individual scale long, pointed apically and fringed nearly to tip. Hair 0(1), 1(20, 17- 24), 2(1), 3(7, 5-8), 4(1), 5(15, 12-17), 14(1). Siphon: Index about 2.1-2.4; integu- ment moderately pigmented, imbricate. Pecten extending to 0.42-0.45, with 13-20 teeth; individual tooth long, pointed, with basal denticles. Hair 1(2, 2- 3): 2(1), 3-5 not visible, 6(1), 7(1), 8(1, 1-2), 9(1). Anal Segment: Saddle moderately pigmented and weakly spiculate; caudal margin with spines of varying length. Gills moderate, about 1.5-2.0 of saddle length, ventral slightly shorter than dorsal. Hair 1(4, 3-5), 2(5, 3-6), 3(2), 4a(3, 2-4), 4b(2, 2-3), 4c(2, 2-3), 4d@2, 2-3), 4e(2, 2-3). EGG. Length about 0.73 mm, width 0.1 mm; exochorion transparent, with cir- cular clear refractile bodies; endochorion dark, with irregular polygonal sculpturing. SYSTEMATICS. Formerly the name sexlineatus was applied to quite unrelated forms, some of which are now recognized as distinct species in other groups of the Sexlineatus Section. In the restricted sense used here, with the diagnostic features mentioned under the group and the species, sexlineatus appears to be a complex of several closely related forms. It is impossible at this time to resolve the complex owing to the paucity or total absence of associated immature stages and even adults from critical areas. The best diagnostic features of the complex are: (1) in the adults, the presence of dark scales on the upper ppn and antealar area; and (2) in the larvae, the presence of a dense vestiture of hairlike spicules on the thorax and abdomen; the aedeagus of the male genitalia may also prove to be distinctive. I have seen samples from populations with these characters from central and southern Mexico, Honduras, Panama, Ecuador, Venezuela and Trinidad. The species has been reported from inter- mediate areas but little reliance can be placed on these records although probably the complex does occur throughout Central America and in Colombia. The description of the species is based on topotypic associated material from Trin- idad. The females and 4 males from Venezuela are so similar to those from Trinidad except for slightly yellowish mesonotal lines that there is no question that they be- long to the same taxon. No significant departure was noted in the limited material of immature stages from Venezuela. From the original description and illustration it is evident that dominicii Rangel and Romero, 1907 from the Rio Anauco is based on this population and it is therefore synonymized here with sexlineatus. The Trinidad and Venezuela populations appear to be confined to the coastal areas and are known from near sea level to moderate elevations in the coastal ranges. The Ecuador population is known only by males and females from the lowlands of the Guayas River basin. These specimens resemble the typical populations except for fewer dark scales on the upper ppn and a tendency toward more extensive light scaling on the tarsi, some specimens showing some white scales on segment 3 of the foretarsus, segments 3 and/or 4 of the midtarsus and segments 4 and/or 5 of the hindtarsus. In the male genitalia, some of the setae of the ninth tergite lobe are slight- ly flattened, approaching the condition characteristic of ecuadoriensis of the Elea- norae Group. Limited material of all stages has been examined from Panama. While the majority of adults agree with the typical populations, 2 show a preponderance of white scales on the upper ppn. I have not been able to detect any significant differences in the immature stages from the typical populations. The 3 females from Honduras, 2 females from Colombia and all adults from Mexi- co differ from the typical populations apparently only in the presence of fewer dark scales cn the upper ppn. In the limited material of immature stages from Mexico no significant differences from typical populations were noted. These populations are found at higher elevations than the typical ones, particularly the ones from Cuerna- Berlin: Subgenus Howardina 91 vaca, Mexico (about 7,000 feet). In the southern portion of the range, populations of sexlineatus are apparently largely or entirely restricted to the rain forest and tropical deciduous forest at low and moderate elevations. This may be the case even in Central America and Mexico where sexlineatus populations are known only from higher elevations where patches of such forests may be present. BIONOMICS. Immature stages of sexlineatus have been collected only in epiphy- tic bromeliads. Females are readily attracted to human bait and Trapido and Galindo (1957) found that in Panama they bite both at ground level and in the forest canopy. Attempts at experimental infection with the yellow fever virus have yielded negative results in Trinidad (Aitken 1960:4). Similar negative results have been reported by Boshell-Manrique and Osorno-Mesa (1944:174,177,178) but it is doubtful that the Species involved was actually sexlineatus. DISTRIBUTION (fig.4). Mexico (southern part), probably Guatemala, Honduras, Nicaragua, possibly Costa Rica, Panama, Colombia (upper Orinoco drainage), Vene- zuela (northern), Trinidad and Ecuador (Guayas River basin) at elevations of 250- 7006 ft. Material examined: 1099 specimens; 28 6, 684 ?, 35 pupae, 352 larvae; 32 individual rearings (1 pupal, 24 larval, 7 incomplete). COLOMBIA. Boyaca: Cuincha, 2 9; Guamales, 2 9; San Isidro, 3 9 (Boshell-Manrique and Osor- no-Mesa 1944:174,177,178). Meta: Retiro, 15-16-19 Aug 1935, 2 9 [UCLA, USNM]. _ ECUADOR. Babahoyo, Aug 1938, F. Campos, 4 6, 18 2 [USNM] ; Balzapamba, Aug 1938, F. Campos, 4 d, 1 2 [USNM], 6 ? (KO 209A-21), 1939 or 1941, H. Hansen, 2 ? (KO 209A-14) [UCLA] . Guayaquil, J. Murdock, 2 6,3 9 [USNM]. HONDURAS. Tegucigalpa, 15 July 1945, 3 9 [USNM]. : MEXICO. Chiapas: San Cristobal (Martini 1935:49). Tuxtla, 23 July 1963, E. Fisher (MF 8), 21 2; Sumidero, 17 Aug 1964, D. Verity, 1 lpd (117-11), 4 L (MEX 117), 1 lpd (118-10); 17 Aug 1964, E. Fisher and D. Verity, 1 9? (MEX 120), 1 9 (MEX 121), 8 9 (MEX 128) [UCLA]. Morelos: Cuernavaca, elev. 7000 ft, 29 July 1961, R. and K. Driesbach, 2 6, 2 9 [USNM]. Veracruz: Cordo- ba, 12 July 1964, E. Fisher and D. Verity, 5 9 (MEX 18), 1 9? (MEX 19), 1 9 (MEX 21); 13 July 1964, E. Fisher and D. Verity, 3 9 (MEX 26), 1 Ip¢ (28-14), 1 Ip? (28-11); 16 July 1964, E. Fisher and D. Verity, 1 L (MEX 38) [UCLA]. NICARAGUA. Villa Somoza, Aug, Sept, Nov 1953, 1 9 (Galindo and Trapido 1957:146). PANAMA. Bocas del Toro: Almirante, 1952-53, 77 A (Trapido and Galindo 1957:122,125). Cocle: El Valle, elev. 2000 ft, 14 Aug 1963, A. Quinonez (PA 533), 1 IP (533-101); Valle de Anton, elev. 2400 ft, 13 Aug 1963, A. Quinonez (PA 504), 3 Ip (504-101,102,108), 1 Ip? (504-104), 1 pd (504-103), 1 IP (504-106) [UCLA]. Tucue, elev. 1000 ft, 12 June-14 Dec 1950, 2 A (Trapido, Galindo and Carpenter 1955:530). Panama: Campana, elev. 2000 ft, 14 June-6 Dec 1950, 24 A (Trapido, Galindo and Carpenter 1955:530). TRINIDAD. Caroni: Brasso Venado, elev. 400 ft, 21 June 1964, R.L. Manuel and R. Martinez, 24 9 (TR 513) [UCLA]. Mayaro: Bush Bush Forest, 9 Dec 1963, TRVL, 2 9 (TR 62); 20 Dec 1963, TRVL, 1 ? (TR 5); 27 Dec 1963, TRVL, 6 2? (TR 21); 30 Dec 1963, TRVL, 2 2 (TR 10); 17 Jan 1964, TRVL, 4 9° (TR 28); 22 Jan 1964, TRVL, 1 2 (TR 40) [UCLA]. Nariva: Archers Estate, 5 Nov 1964, A. Guerra, 1 9? (TR 819) [UCLA]. Biche, elev, 250 ft, 9 July 1964, F. Powd- har, 3 9 (TR 563) [UCLA]. Charuma Forest, elev. 500 ft, 27 Aug 1964, A. Guerra, 24 9 (TR 644); 8 Oct 1964, A. Guerra, 3 2? (TR 761) [UCLA]. St. Andrew: Caratal Road, elev. 250 ft, 3 Dec 1964, F. Powdhar, 1 ? (TR 866) [UCLA]. Coryal, elev. 250 ft, 18 June 1964, A. Guerra, 29(TR 498) [UCLA]. Mt. Harris, 23-31 July 1924, C.L. Withycombe, 1 9 [USNM]; elev. 250 ft, 16 July 1964, F. Powdhar, 16 9? (TR 573) [UCLA] . Nestor Village, elev. 100 ft, 12 June 1965, A. Guerra, 1 2 (TR 484) [UCLA]. Tamana, 1945, 1 9 [USNM];Mt. Tamana, elev. 250 ft, 19 June 1964, A. Guerra, 43 ? (TR 512); 20 Aug 1964, A. Guerra, 1 9? (TR 619); elev. 1000 ft, 23 Aug 1964, R.L. Manuel and R. Martinez, 5 2? (TR 620) [UCLA]. Valencia, Cumaca, elev. 1000 ft, 3 Sept 1964, A. Guerra, 2 9 (TR 654); El Quemado Road, elev. 250 ft, 3 July 1964, F. Powdhar, 1 9 (TR 547) [UCLA] . St. David: Sans Souci, elev. 250 ft, 5 Mar 1964, A. Guerra, 1 9 (TR 151) [UCLA]. St. 92 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 George: Aripo Valley, elev. 500 ft, 16 Apr 1964, A. Guerra, 1 2 (TR 330) [UCLA] . Blanchisseuse, 2 Aug 1945, 18 9; 2 Sept 1945, 1 9 [USNM]; 1 9 (KO 209A-36) [UCLA] ; road to Blanchisseuse, 2 Mar 1945, 2 9 [USNM]. Guanapo Valley, elev. 1500 ft, 17 Apr 1964, A. Guerra, 1 ? (TR 339) [UCLA]. Heights of Guanapo, elev. 800 ft, 26 Mar 1964, A. Guerra, 1 9 (TR 259); elev. 1000 ft, 22 Apr 1965, A. Guerra, 63 2 (TR 1129) [UCLA]. La Laja Road, elev. 1000 ft, 11 Mar 1965, A. Guerra, 44 ? (TR 1036) [UCLA]. La Laja Trace, 11 Mar 1965, 2 Ipd (1041-22,41), 1 Ip? (1041- 18), 2 lp (1041-21,23), 2 P, 33 L, 13 1 [UCLA]. Las Lapas Trace, elev. 2000 ft, 3 Apr 1964, A. Guerra, 12 9 (TR 281) [UCLA]. Mt. Beck, elev. 1000 ft, 29 Apr 1965, A. Guerra, 1 9 (TR 1132) [UCLA]. Port of Spain, 26 Aug 1945, 1 @ [USNM]. Simla, 25 Mar 1965, M.J. Moody, 6 ° (TRM 15) [UCLA]. Springhill Estate, elev. 1200 ft, 25 Mar 1965, A. Guerra and M.J. Moody (TRM 12), 2 Ip? (12-50,52), 10 L; 25 Mar 1965, A. Guerra, 2 lp? (1056-11,14), 1 P,5 L,11(TR 1056), 3 Ip? (1063-31,81,82), 6 2, 86 L(TR 1063), 2 Ipd (1064-11,12), 1 9, 3 L(TR 1064) [UCLA]. Verdant Vale, elev. 1000 ft, 10 Sept 1964, A. Guerra, 5 9? (TR 686); elev. 500 ft, 12 Nov 1964, A. Guerra, 21 2(TR 831); 9 Apr 1965, elev. 1200 ft, A. Guerra, 3 ?, 132 L(TR 1098), 1 Ipd (1099-33), 3 Ip? (1099-11 21,22), 3 IP (1099-12,31,32), 34 L [UCLA]. County not specified: June 1905, A. Busck, 15 9; A. Busck, 7 9; F.W. Urich, 1 9 [USNM]. : VENEZUELA. Anzoategui: Puerto La Cruz, Sept 1944, W.H.W. Komp, 1 ? (KO 209A-15), 2 6 (KO 209A-20); 26 Sept 1944, W.H.W. Komp, 10 ? (KO 209A-24), 23 2 (KO 209A-27), 24A (KO 209A-35) [UCLA]. Aragua: Guamitas, elev. 1200 m, 27 Jan 1927, M.N. Tovar, 3 2; 24 Aug 1927, M.N. Tovar, 136 9; 27 Aug 1927, M.N. Tovar, 1 ? [USNM]. La Providencia (Cova Garcia, Sutil and Rausseo 1966b:74). Maracay, 2 July 1927, M.N. Tovar, 2 9;M.N. Tovar, 1 9 [USNM]. Rancho Grande, 2 Nov 1962, I. Ortiz C., 7 9? [USNM]. Turmero (Cova Garcia, Sutil and Rausseo 1966b:74). Distrito Federal: Caracas, 21 July 1938, 1 6;P.J. Anduze, 1 6, 1 9 [USNM]; Hacienda El Paraiso, 9 (Rangel and Romero-Sierra 1907). Carayaca (Cova Garcia, Sutil and Rausseo 1966b: 74). Miranda: El Hatillo (Cova Garcia, Sutil and Rausseo 1966b:74). Monagas: Guacharo, F.M. Root, 5 2?(VZR 267) [UCLA]. Whitmorei Group FEMALES. Head: Median light line on vertex with moderately broad scales. Pro- boscis subequal to forefemur in length. Torus with an almost complete ring of white scales broken only dorsally. Thorax: Mesonotal light lines usually white to silvery; inner dorsocentral line complete; outer dorsocentral line shorter, not reaching ante- rior margin, diverging outwards anteriorly; lateral marginal line broken into narrow prescutal, antealar and supraalar patches; scales on anterior promontory white. An- tealar area above paratergite with dark scales. Paratergite and ppn with restricted light patch confined to posterior half; upper stp patch narrow, elongate. Legs: Distal white spot on anterior surface of midfemur present, absent on hindfemur. Tibiae predominantly dark, with light lines or scattered scales. Wing: Veins entirely dark scaled. Abdomen: Basomedian tergal light markings absent. MALES. Palpus predominantly dark, white scales restricted to base of segments 4 and 5. MALE GENITALIA. Lobe of tergite [IX prominent, with a few (5-7) strong or many (about 12) weak bristles. Sidepieces separated at base by a distance greater than width of aedeagus; tergomesal and/or sternomesal margins with more or less distinct median lobes. Claspette completely absent. Aedeagus moderate to long, shorter than or subequal to proctiger; apex deeply emarginate, with distinct apical horns. PUPA (whitmorei only). Cephalothorax: Trumpets short, index about 3.0, not expanded apically; hair 10-C branched, 12-C single or double. Abdomen: Hair 5-IV-V moderate, not exceeding apex of second tergite following; hair 9-VII moderate, Berlin: Subgenus Howardina 93 branched, exceeding lateral angle of tergite. Paddle: Moderate; apex rounded. LARVA (whitmorei only). Head: Hairs 4-7-C moderately pigmented, all at least 5-branched; hair 15-C moderate, exceeding apex of mental plate. Thorax: Squarish in outline; integument strongly spiculate, spicules hairlike, simple. Abdomen: Integ- ument strongly uniformly spiculate. Hair 8-VII slightly cephalodorsad of 9-VII. Si- phon: integument distinctly spiculate; acus distinct, attached. Anal Segment: Hair 3- X single; ventral brush with 5 pairs of hairs. DISCUSSION. The Whitmorei Group of 2 species is the subgenus Soperia of au- thors, which is based on a misidentified species (see discussion under subgenus How- ardina). This group is a very distinct one in adult ornamentation and male genitalia but is obviously related to other groups in the Sexlineatus Section. Its closest affini- ties seem to be with the Sexlineatus and Allotecnon Groups. The immature stages of whitmorei are less differentiated from other groups of the Sexlineatus Section than are those of walkeri from other groups of the Walkeri Section. The Whitmorei Group is known at present only from Colombia, Venezuela, Bo- livia and probably Argentina (fig.5). Although whitmorei and pseudodominicii have been found together in Meta, Colombia, the 2 species may be largely allopatric, whit- morei occupying the northern part of the range of the group. A. pseudodominicii penetrates into the Amazon basin at an elevation of less than 2,000 feet in Chimore, Bolivia and apparently into the La Plata basin at moderate elevations in northern Ar- gentina. It is also possible that this group is represented on the eastern slope of the Andes in Ecuador (Levi-Castillo 1953:40). 22. Aedes (Howardina) whitmorei Dunn Figs.5 ,9,45 46 1918.Aedes whitmorei Dunn, 1918:128-130. TYPE: Lectotype 2, near emerald mines, Muzo, Co- lombia, 1916, E.R. Whitmore [USNM; selection of Stone and Knight 1956:226]. Aedes (Howardina) whitmorei of Bonne and Bonne-Wepster (1925:375); Dyar (1928:230); Ed- wards (1932:156); Horsfall (1955 :471). Aedes (Soperia) whitmorei of Lane (1939:98); Anduze (1941:14; 1943:193; 1947:357); Lane (1953:701-703); Reyes (1955:74,89); Stone and Knight (1956:226); Stone, Knight and Starcke (1959:173); Belkin, Schick and Heinemann (1965:9); Forattini (1965 :405); Cova Garcia, Sutil and Rausseo (1966a:57,275; 1966b:80-81,141 339). Aedes whitmorei of Antunes (1937b:77); Senevet and Quievreux (1941:258); Boshell-Manrique and Osorno-Mesa (1944:173,174,175,177,178). Aedes (Soperia) dominicii of Komp (1936a:63; 1936b:73-74). Aedes dominicii in part of Bates (1944:165 166; 1945:23; 1949:16,61,320); Bugher, Boshell-Man- rique et al (1944:33,37). FEMALE. Wing: 2.7 mm. Proboscis: 1.98 mm. Forefemur: 1.98 mm. Abdomen: about 2.4 mm. Head: Light decumbent scales on vertex silvery white. Erect scales light on occiput, darkened on sides. Palpus about 0.16 of proboscis, with silvery white scales on apex of segment 4. Torus with silvery white scales. Thorax (fig.9): Mesonotal light lines white; inner dorsocentral line very narrow; outer dorsocentral line narrow but with a distinct broad silvery spot of broad scales at anterior end in fossa; lateral marginal line broken into lateral prescutal line, a small antealar and a larger supraalar patches, lateral prescutal nearly continuous but apparently broken at level of humeral bristle. Median scutellar lobe laterally with broad dark scales and 94 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 with median broad silvery white scales from base to apex; lateral lobes with a few narrow silvery white scales. Scales on paratergite broad, silvery. Pleural scales broad and silvery; lower ppn patch adjoining upper ssp; other patches as in the group. Legs: Forefemur predominantly dark on both sides except for narrow white longitudinal line posteriorly; midfemur dark anteriorly except for a silvery spot at distal 0.67, proximal 0.5 of posterior surface white, rest dark; hindfemur predominantly white anteriorly to 0.67, distal part dark, proximal 0.5 of posterior surface white, distal dark. Tibiae usually dark, a few whitish scales at base. Foretarsus and midtarsus with short incomplete white rings on segments 1 ,2; hindtarsus with moderately broad ones on 1-3. Haltere: Stem pale, knob predominantly dark, fringe scales whitish. Abdo- men: Basomedian tergal pure white scales evident on segments V-VII. Sternites with broad basal V-shaped pure white patches; apical light scales inconspicuous. MALE. Wing: 2.5 mm. Proboscis: 2.34 mm. Forefemur: 1.89 mm. Essentially sim- ilar to female in color; only apparent difference is a small ventromedian white spot on the proboscis. MALE GENITALIA (fig.45). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with 6,7 moderately strong bristles. Sidepiece rather short, length about 2.0 of median width; tergomesal margin with a row of unpigmented leaflike setae, sternomesal lobe with a dense tuft of pigmented leaflike setae on an elevated flap. Clasper about 0.5 of sidepiece, with spicules in basomesal area; spini- form moderately long, about 0.35 of clasper length. Aedeagus narrow, subequal to proctiger, apex deeply emarginate, with distinct horns curved laterally. PUPA (fig.45). Abdomen: 2.86 mm. Trumpet: 0.32 mm; index about 3.0. Paddle: 0.67 mm. Diagnostic characters as in the key. Chaetotaxy as given below, hairs mod- erately pigmented. Cephalothorax: Mesonotum lightly pigmented between trumpets, lateral areas slightly darker; metanotum uniformly moderately pigmented. Trumpets short, strongly pigmented and contrasting. Hair 1(1), 2(1), 3(1), 4(3, 2-4), 5(2), 6(1), 7(2), 8(1, 1-3), 9C1, 1-2), 10(4, 2-5), 11(1), 12(2, 1-2). Abdomen: Tergite I uniform- ly lightly pigmented, except for usual membranous areas; rest of segments imbricate and lightly pigmented, progressively lighter caudad. Segment I: hair 1(7-12 primary branches, each with few secondary ones), 2(1), 3(1), 4(2, 2-3), 5(1, 1-3), 6(1), 7(1), 9(1), 10(1, present in a few), 11(2, 1-4, present in a few). Segment II: hair 0(1), 1(4, 2-5), 2(1), 31), 4(@2, 2-3), 5(1, 1-2), 6(1), 7(1, 1-2), 8(1), 9(1), 10(1). Segment IIT: hair OC] ), 1G, 2-3), 201), 3€2), 4C1, 1I-2), 5(1, 1-2), 6(1), 7@, 1-3); 8(1, 1-2), 901), 10(1), 11€1), 141). Segment IV: hair 0(1), 1(2, 2-4), 2(1), 3(2, 2-3), 4(1), 5(2), 6(1), 7(2, 2-4), 8(1, 1-2), 9(1), 101), 11(1), 14(1). Segment V: hair 0(1), 1(2, 2-3), 2(1), 3(1), 4(2, 1-4), 5(1), 6(1), 7(2, 2-5), 8(1, 1-2), 9(1), 10(1), 1101), 14(1). Segment VI: hair O(1), 1€1, 1-3), 21), 31), 4(1, 1-2), 5(2), 6(1), 71, 1-3), 8(1, 1-2), 9(1), 10Q1, 1-2), 11(1), 141). Segment VII: hair 0(1), 1(1, 1-2), 2(1), 3(1), 4(1), 5(1), 6Q, 2-3), 7(1), 8(3, 3-4), 9(5, 4-6), 10(1), 11€1), 14€1). Segment VIII: hair 0(1), 4(1), 9(10, 8- 11), 141, 1-2). Paddle: Uniformly lightly pigmented, midrib slightly darker; external margin serrated. Male genital lobe extending to 0.5 of paddle, female genital lobe to 0.39. LARVA (fig.46). Head: 0.84 mm. Siphon: 0.81 mm. Anal Saddle: about 0.3 mm. Diagnostic characters as in the key. Head: Width subequal to length; pigmentation moderate except around mouthparts where it is deeper. Mental plate with a strong median tooth and 11 distinct teeth on each side, often a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 4(15, 10-16), 5(6, 5-8), 6(7, 6-7), 7(14, 11-17), 8(3, 3-6), 9(11, 8-14), 10(1), 11(20, 16-20), 12(5, 4-7), 13(1), 14(14, 11-14), 15(3, 2-3). An- tenna: Length about 0.32 of head; shaft rather uniformly pigmented and with a few Berlin: Subgenus Howardina 95 scattered spicules; hair 1(3, 2-3), other single. Thorax: Stellate hairs with many un- equal, weakly barbed branches; long hairs attenuate apically and weakly barbed. Pro- thorax: hair 0(19, 19-23), 1(5, 4-6), 2(1), 3(21, 13-23), 4(1), 5(1), 6(1), 7(2), 8(@20, 16-20), 9(14, 14-17), 10(€1), 11(€1), 12€1), 13(22, 20-25), 14(34-51). Mesothorax: hair 1(14, 13-20), 2(1), 3(1), 41), 5(1), 6(5, 4-6), 7(1), 8(6, 6-7), 9(5, 4-6), 10(1), 11(2, 1-3), 12(1), 13(17, 10-19), 14(16, 16-21). Metathorax: hair 1(25, 20-27), 2(1), 3(2, 1-2), 418, 17-19), 5(18, 17-23), 6(1), 7(5, 4-6), 8(18, 16-22), 9(4, 3-6), 10(1), 11(2, 1-2), 12(1), 13(22, 19-24). Abdomen: Integument and stellate hairs as on tho- rax. Segment I: hair 1(23, 20-25), 2(20, 20-25), 3(1), 4(1), 5(16, 14-23), 6(2, 2-3), 7(1), 9(8, 7-8), 10(1), 11018, 15-20), 13(22, 22-25). Segment II: hair 0(1), 1(21, 20- 24), 2(22, 18-25), 3(1), 4(1), 5(24, 18-24), 6(2, 2-3), 7(10, 9-10), 8(1), 9(14, 9-15), 101), 111), 121), 13@20, 19-24). Segment III: hair 0(1), 1(22, 19-25), 2(21, 18- 27), 3(1), 4(1), 5(19, 17-26), 6(2), 7(17, 14-20), 8(1), 9(16, 14-21), 10(1), 1101), 12(1), 13(@20, 16-26), 14(1). Segment IV: hair 0(1), 1(20, 17-24), 2(22, 20-22), 3(1), 4(1), 5(22, 18-25), 6(2), 7(15, 9-18), 8(1), 9(19, 18-23), 10(1), 11(1), 121), 1318, 14-20), 14(1). Segment V: hair 0(1), 1(22, 16-23), 2(19, 17-23), 3(1), 4(1), 5(20, 18- 23), 6(2), 7(15, 12-18), 8(1), 9(19, 18-24), 10(1), 11(1), 12(1), 1319, 16-22), 14(1). Segment VI: hair 0(1), 1(19, 19-23), 2(19, 18-23), 3(1), 4(1), 5(21, 18-22), 6(1), 7(11, 10-18), 8(1), 9(20, 17-21), 10(1), 111), 12(1), 13(17, 13-18), 14€1, 1-2). Seg- ment VII: hair 0(1), 1(20, 18-21), 2(18, 16-22), 3(1), 4(1), 5(16, 16-22), 6(10, 10- 13), 71), 810, 8-11), 9(16, 11-18), 10(1), 11(1), 12(1), 13(11, 10-14), 14(@2, 1-2). Segment VIIT: Comb scales usually 12(9-12) in number, in a single row; individual scale long, pointed apically and fringed almost to tip. Hair 0(1), 1(28, 22-28), 2(1), 3(8, 7-9), 4€1), 5(15, 10-19), 14(2, 2-3). Siphon: Index about 3.0, distinctly swollen in middle; integument uniformly moderately to strongly pigmented, with darker basal ring, uniformly spiculate. Pecten teeth usually 16(14-20) in number, extending to 0.4 before hair 1-S. Hair 1(3, 2-3), 2(1), 3-5 not visible, 6(1), 7(1), 8(1, 1-2), 9(1). Anal Segment: Saddle strongly pigmented, with darker basal ring; integument uni- formly covered with spicules; caudal margin with spines of varying length; gills sub- equal, long, more than 2.0 of saddle length. Hair 1(2), 2(4, 3-4), 3(1), 4a(2), 4b(2), 4c(2, 1-2), 4d(2, 1-2), 4e(1, 1-2). SYSTEMATICS. A. whitmorei is a very strikingly differentiated species errone- ously synonymized with dominicii by Komp (1936a:53); the latter as shown here is undoubtedly sexlineatus (see). Its only close relative is pseudodominicii, whose im- mature stages are unknown. In ornamentation of the adults and in male genitalia the 2 species are readily separated by the features mentioned in the diagnoses and keys. A. whitmorei is known at present from both sides of the Cordillera Oriental in Co- lombia and from the Cordillera de la Costa in northcentral Venezuela; it probably occurs also in the intermediate areas in the Cordillera de Merida in western Venezue- la. On the eastern slope of the Cordillera Oriental whitmorei is sympatric with pseu- dodominicii whose distribution extends on down to northern Argentina along the eastern slopes of the Andes. BIONOMICS. Immature stages of whitmorei have been collected in epiphytic bro- meliads at elevations of about 3,000 feet. Dunn’s (1918:130) record from ‘“‘small heavily shaded pools of clear but apparently stagnant water” is probably erroneous. According to Bates (1949:16,61), whitmorei (as dominicii) is a relatively common diurnal forest mosquito in the region of Villavicencio in Colombia; the females bite both at ground level and in tree stations; and males are often observed hovering over men and animals in the forest. Experimental infections of this species (also as domi- nicii) with yellow fever virus have been unsuccessful (Bugher, Boshell-Manrique et al 96 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 1944:33,37). DISTRIBUTION (fig.5). Colombia (upper Orinoco and Magdalena drainages) and Venezuela (Cordillera de Merida and Cordillera de la Costa) at moderate elevations. Material examined: 104 specimens; 32 6, 43 9, 7 pupae, 22 larvae; 7 individual larval rearings. COLOMBIA. Boyaca: Las Animas, 500 yds from mine, 17 Aug 1916, 6 ? [USNM] . Muzo, em- erald mines, E.R. Whitmore, 1 9; 16 Aug 1916, E.R. Whitmore, 10 , type series [USNM]. Guamales, 51 9; Tamborredondo, 1 9; Cuincha, 162 2; San Isidro, 37 2 (Boshell-Manrique and Osorno-Mesa 1944:174,175,177,178). Meta: Palucho, 3 6, 2 2? [USNM]. Restrepo, 17 July 1935, W.H.W. Komp, 1 6, 1 2 (KO 203A-10); 20 Aug 1935, W.H.W. Komp, 1 6 (KO 209A-2) [UCLA]; Aug 1935, W.H.W. Komp, 2 6, 3 2; 9 Sept 1935, W.H.W. Komp, 1 6,2 ?, 101 [USNM], 2 6, 3 2 (KO 209A- 28) [UCLA]; 1935, W.H.W. Komp, 1 6 (KO 209A-40), i 6 (KO 209A-42), 1 d (KO 209A-44) [UCLA]; July 1935, 2 6; Sept 1935, 2 9; 1935, 2 6; 4 June 1940, 1 6; 2 d [USNM]; Retiro, 22 Aug 1935, W.H.W. Komp, 5 6; 15-16-19 Aug 1935, 2 ? [USNM]. Villavicencio, Buena Vista, elev. 3000 ft, 9 July 1965, E. Osorno-Mesa et al (COB 68), 1 Ipd (68-30), 2 L [UCLA] ; 4 June 1942, W.H.W. Komp, 2 6 [USNM]; Forzosa, 25 Oct 1943, M. Bates, 1 ? [USNM] ; Quebrada Colorado, elev. 2300 ft, 9 July 1965, E. Osorno-Mesa et al (COB 67), 4 Ipd (67-20,22,23 125), 2 Ip? (67-21, 24),3 L [UCLA]. VENEZUELA. Aragua: Ocumare de la Costa, 5 July 1927, M.N. Tovar, 5 ? [USNM]. Rancho Grande, 2 Nov 1942, I. Ortiz C., 3 9 [USNM]. Distrito Federal: Caracas (Anduze 1943:193; Cova Garcia, Sutil and Rausseo 1966b:80). Cementerio (Cova Garcia, Sutil and Rausseo 1966b:81). Merida: Bailadores (Cova Garcia, Sutil and Rausseo 1966b:81). Tovar (Anduze 1947:357). Tachira: Jaurequi and Uribante (Anduze 1947:357). : 23. Aedes (Howardina) pseudodominicii Komp Figs.5 9,58 1936.Aedes pseudodominicii Komp, 1936a:75. TYPE: Holotype 6, Restrepo, Meta, Colombia, Aug-Sept 1935, E. Osorno-Mesa [NE]. 1955.Aedes (H.) delpontei Martinez and Prosen, 1955:35-40. TYPE: Holotype 6, Chapare, Cocha- bamba, Bolivia, elev. 6000 ft, Jan 1949, Feb 1952 or Nov 1953, Martinez and Prosen [MEPRA]. NEW SYNONYMY. Aedes (Soperia) pseudodominicii of Komp (1936b:63; 1956:37). Aedes (Soperia) dominicii of Antunes (1937a:16; 1937b:76-77); Lane (1939:98); Senevet and Quievreux (1941:258); Anduze (1943:192-193); Boshell-Manrique and Osorno-Mesa (1944:173, 174,175,177,178); Levi-Castillo (1952:555; 1953:40); Lane (1953:700-701); Reyes (1955:73, 89); Stone , Knight and Starcke (1959:173); Forattini (1965: 405). Aedes (Soperia) dominicii in part of Bates (1944:165 166; 1945:23; 1949:16,61,320); Bugher, Bo- shell-Manrique, et al (1944:33,37). Aedes (Howardina) dominicii of Horsfall (1955:471). Aedes (H.) delpontei of Martinez and Prosen (1959:603,605); Stone, Knight and Starcke (1959: 174); Prosen, Carcavallo and Martinez (1964:104,119). FEMALE. Wing: 3.42 mm. Proboscis: 2.61 mm. Forefemur: 2.61 mm. Abdomen: about 3.24 mm. Similar to whitmorei but usually slightly larger in size and differing in the following. Head: Light decumbent scales white. Erect scales on occiput and sides whitish. Palpus about 0.17 of proboscis, with a few mesal and apical white scales on segment 4. Torus scales white. Thorax (fig.9): Mesonotal light lines white, rarely faintly creamy; inner and outer dorsocentral line without silvery spot in fossal area. Scutellar light scales white. Pleural scale patches white and distributed as in Berlin: Subgenus Howardina 97 whitmorei. Legs: Forefemur predominantly auburn except for a few basal white scales anteriorly and a narrow longitudinal line posteriorly; midfemur predominantly auburn except for a white spot at distal 0.67 anteriorly and a mesal white line to middle posteriorly; hindfemur predominantly white on both sides, rest mostly au- burn; midfemur and hindfemur with a short mesal patch merging with knee spot. Foretibia predominantly auburn, with a few scattered white scales; midtibia and hindtibia with a continuous white line, rest auburn. Foretarsus with a few white scales on segment 1; midtarsus with short incomplete white ring on 1; hindtarsus with broader white rings on segments 1,2. Haltere: Stem brownish, knob predominantly auburn, with a few white fringe scales. Abdomen: Basal sternal light scales white, apical white scales apparent to segment V, rest auburn. MALE. Wing: 2.52 mm. Proboscis: 2.34 mm. Forefemur: 1.9 mm. Essentially sim- ilar to female in color. Proboscis completely dark. MALE GENITALIA (fig.58). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with many (about 12) narrow bristles. Sidepiece rather short, broader at the middle, tergomesal area with few and sternomesal area with numerous narrow setae. Clasper about 0.6 of sidepiece, stout, curved; spiniform short. Aedea- gus shorter than proctiger; dorsal sclerotization deeply emarginate, apical horns dis- tinct, curved upwards and dorsally. PUPA, LARVA unknown. SYSTEMATICS. A. pseudodominicii is a distinct, clearly marked species errone- ously synonymized by Antunes (1937a:16) with dominicii; the latter is actually sex- lineatus (see). On the basis of ornamentation of the adults (only stages known), pseu- dodominicii is evidently closely related to whitmorei, which has also been confused (Komp 1936a:53; 1936b:73-74) with dominicii. The 2 species are easily distinguished by the diagnostic features given above. A. pseudodominicii apparently occurs at mod- erate elevations only along the eastern slopes of the Andes, from the Cordillera Ori- ental in Colombia, where it is sympatric with whitmorei, all the way down to north- ern Argentina. It penetrates the upper reaches of the 3 main eastern drainage sys- tems of South America, the Orinoco, Amazon and La Plata. Although I have not seen any of the type material of delpontei Martinez and Prosen, 1955, it is evident that this population from Bolivia is conspecific with the typical population from Colom- bia as the original description of delpontei obviously matches in all pertinent details the diagnostic features of pseudodominicii as do 2 females collected by Torres-Munez at the type locality of delpontei. I have not seen the female paratype of delpontei from Argentina but see no reason to doubt its conspecificity with the material from Bolivia and Colombia. BIONOMICS. The immature stages of pseudodominicii are unknown but undoubt- edly will be found in epiphytic and/or terrestrial bromeliads. It is very likely that the reports of Bates (1949) and Bugher and Boshell-Manrique et al (1944) mentioned un- der whitmorei apply at least in part to pseudodominicii since at that time both spe- cies were probably identified as dominicii. DISTRIBUTION (fig.5). Colombia (upper Orinoco and Magdalena drainages), Bo- livia (upper Amazon drainage) and Argentina (upper La Plata drainage) at moderate elevations. Material examined: 20 specimens; 10 6, 10 2; no individual rearings. ARGENTINA. Salta: Campamento San Pedro S.0.C., Departamento San Martin, 1 ? paratype of delpontei (Martinez and Prosen 1955:40). BOLIVIA. Cochabamba: Chapare, Chimore, 29 June 1944, Torres-Munoz, 1 ?; 29 Apr 1944, 1 9 [USNM]. Za Paz: Nor Yungas, A (Prosen, Carcavallo and Martinez 1964:104). COLOMBIA. Boyaca: La Bodega, 22 ?;Guamales, 58 2; Tamborredondo, 33 2; Cuincha, 19 9; 98 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 San Isidro, 15 ? (Boshell-Manrique and Osorno-Mesa 1944:173,174,175,177,178). Meta: Palucho, 2 6 [USNM]. Restrepo, 26 July 1935, W.H.W. Komp, 1 ? (KO 209A-39); 20 Aug 1935, W.H.W. Komp, 4 6 (KO 209A-2) [UCLA]; 7 Sept 1935, E. Osorno-Mesa, 1 6; 1935, W.H.W. Komp, 1 d; July 1935, 1 9 [USNM]; Retiro, 26 July 1935, W.H.W. Komp, 3 ? (KO 209A-31) [UCLA] ; 15-19 Aug 1935, 16, 2 9 [USNM]. Villavicencio, Buena Vista, 4 June 1942, W.H.W. Komp, 1 d [USNM]. Allotecnon Group FEMALES. Head: Median light line on vertex with narrow scales. Proboscis dis- tinctly longer than forefemur. Torus with a restricted mesal scale patch. Thorax: Mesonotal light lines golden; inner and outer dorsocentral lines complete, subparallel; lateral marginal line continuous; scales on anterior promontory creamy to golden. Antealar area above paratergite with golden scales. Paratergite with a moderate scale patch extending entire sclerite length. Upper ppn entirely covered with light scales; upper stp patch narrow, elongate. Legs: Distal white spot on anterior surface of mid- femur and hindfemur present. Tibiae completely dark. Wing: Veins entirely dark scaled. Abdomen: Basomedian tergal light markings present on III-VII. MALE (allotecnon only). Palpus with basomedian light scales on segments 4 and a MALE GENITALIA (allotecnon only). Lobe of tergite IX not prominent, with 3,4 strong bristles. Sidepieces separated at base by a distance less than width of aedeagus; without any indication of lobes on tergomesal or sternomesal margins. Claspette represented by a conical tubercle. Aedeagus shorter than proctiger; apex shallowly emarginate, lateral lobes inconspicuous. PUPAE. Cephalothorax: Trumpet moderate, index about 6.5, not expanded api- cally; hairs 10,12-C usually single or double. Abdomen: Hair 5-IV-V long, usually ex- ceeding apex of second tergite following; hair 9-VII short, slightly larger than 6, usu- ally not exceeding lateral angle of tergite. Paddle: Moderate to long, usually slightly produced on apex. LARVAE. Head: Hair 4-C weak, lightly pigmented, at least 4-branched; 5-C sin- gle, moderate, strongly pigmented; 6-C heavy, strongly pigmented, double; 7-C mod- erate, double or triple; 15-C very short, not extending cephalad of base of mental plate. Thorax: Quadrate in outline; integument glabrous. Abdomen: Integument covered with small spinelike spicules, restricted to dorsal and ventral surfaces (only ventral in guerrero). Hair 8-VII caudolaterad of 9-VII. Siphon: Integument imbricate; acus distinct, separate. Anal Segment: Hair 3-X double; ventral brush with 5 pairs of hairs. DISCUSSION. The Allotecnon Group of 3 species is readily recognized by the unique pleural scale pattern of the adults (particularly the long narrow upper stp scale patch) and the short, spinelike spicules on the abdominal segments of the lar- vae. The group is known only in Central America and is sympatric with the Quadri- vittatus Group from southern Mexico to central Panama (figs.5,6). A. allotecnon, the only widespread species, occupies primarily the central and southern parts of the range of the group but is also known from Chiapas and the southern end of the Sierra Madre Oriental in Mexico; guerrero is apparently endemic to the Sierra Madre del Sur in Mexico; and guatemala is known only from 2 collections on the Pacific slope of central Guatemala. The 3 species appear to be allopatric. Berlin: Subgenus Howardina 99 24. Aedes (Howardina) allotecnon Kumm, Komp and Ruiz Figs.6,9 47,48 1940.Aedes (Howardina) allotecnon Kumm, Komp and Ruiz, 1940:417. TYPE: Lectotype d with associated larval skin, Poas Volcano, Costa Rica, elev. 9000 ft, 14 Apr 1933, H.W. Kumm [USNM; selection of Stone and Knight 1956:214]. : Aedes (Howardina) allotecnon of Kumm and Komp (1941:18-22); Kumm and Zuniga (1942:404); Trapido, Galindo and Carpenter (1955:530,534); Stone and Knight (1956:213); Vargas (1956: 22); Stone, Knight and Starcke (1959:173); Belkin, Schick and Heinemann (1965:12); Forattini (1965:399); Stone (1967:208). FEMALE. Wing: 3.5 mm. Proboscis: 2.71 mm. Forefemur: 2.1 mm. Abdomen: about 3.24 mm. Head: Light decumbent scales on median stripe and short orbital line creamy; broad light scales on sides and venter whitish. Erect scales on occiput dark. Palpus about 0.19 of proboscis, with white scales on apex and mesal side of segment 4. Thorax (fig.9): Inner and outer dorsocentral lines creamy to pale golden, com- plete; lateral marginal line short, extending from scutal angle to wingbase, appearing broad due to presence of creamy scales in antealar area above paratergite. Median scutellar lobe with moderately broad dark scales laterally and median creamy to golden narrow scales from base to apex; lateral lobes with a few narrow creamy scales. Paratergite with broad white scales. Apn with predominantly broad white scales, some anterior ones narrow, creamy. Ppn with large upper patch of narrow creamy scales and small lower patch of broad white scales; psp without scales; other scale patches silvery white and distributed as in group. Legs: Forefemur predomi- nantly dark anteriorly except for basal creamy scales, posterior surface mostly cream- y to middle and with a narrow light line bordered on sides by dark scales extending distally; midfemur mostly dark anteriorly except for basal creamy scales and a sil- very spot at distal 0.67, mainly creamy posteriorly, dark scales restricted to distal half; hindfemur creamy anteriorly to 0.67 and dark distally except for a silvery spot at 0.75, creamy posteriorly to proximal 0.5, rest dark. Foretarsus with basal white scales on segment 1; midtarsus with short, incomplete white rings on 1,2; hindtarsus with moderate white rings on 1-3. Haltere: Stem pale, knob mainly dark, a few fringe scales whitish. Abdomen: Basomedian tergal light scales white from segment III distad. Sternites with broad basal V-shaped patches, creamy at base, silvery on sides; median apical light scales evident to segment IV. MALE. Wing: 2.62 mm. Proboscis: 2.19 mm. Forefemur: 1 q71 mm. Essentially similar to female in color. Mesal light scale patch on torus absent. MALE GENITALIA (fig.47). As figured; diagnostic characters as in the key. Ninth tergite lobe not prominent, with 3,4 strong setae. Sidepiece length 3.0 of median width. Clasper about 0.6 of sidepiece, with 2 preapical hairs; spiniform short, about 0.25 of clasper. Aedeagus shorter than proctiger; dorsal sclerotization shallowly emar- ginate, lateral lobes inconspicuous. PUPA (fig.47). Abdomen: 3.08 mm. Trumpet: 0.43 mm; index 6.5. Paddle: 0.7 mm. Diagnostic characters as in the key. Chaetotaxy as given below; hairs moderate- ly to darkly pigmented. Cephalothorax: Mesonotum moderately pigmented between trumpets, lateral areas and metanotum slightly darker; trumpets contrasting. Hair 1(1), 21, 1-2), 3(1, 1-2), 4(2, 2-4), 5(2), 6(1, 1-2), 7(2), 8C1, 1-3), 91), 10€1, 1-4), 11(1), 12(2, 1-2). Abdomen: Tergite I deeply pigmented, except for usual membra- nous areas; tergites II and III deeply pigmented, other segments lighter. Segment I: 100 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 hair 1(5-13 primary branches, each with many secondary ones), 2(1, 1-2), 3(1, 1-2), 4(3,.2-5), 5(1; 1-2), 6C1, 1-2), 7C1, 1-2), 901), alveoli of 10,11 present in some. Seg- ment II: hair 0(1), 1(3, 2-6), 2(1), 31, 1-2), 4(3, 2-5), 5C1), 6C1, 1-2), 7C1, 1-2), 81, present in a few), 9(1). Segment III: hair 0(1), 1(3, 2-4), 2(1), 3(1), 4(1, 1-2), 5(1), 6(1), 7(2, 1-4), 81, 1-3), 91) 10(1), 11(1), 141). Segment IV: hair 0(1), 1(@, 2-3), 2(1), 3(2, 1-4), 41); SQ, 1-3), 60) 703/14). 80 1-2), 9G), 10), FD, 14C1): Segment V: hair 0(1), 1(2, 1-3), 2(1), 3(1), 4@, 1-3), 5(2, 1-2), 6(1), 7@, 2-4), 8(1, 1-2), 9(1), 101), 11€1), 14(1). Segment VI: hair 0(1), 1(1, 1-2), 2(1), 31), 4(1), 5(1, 1-2), 6€1), 7C1, 1-2), 8(1,.1-2), 901), 1001), 1101), 14C1). Segment VII: hair 0(1), ty, 207, 3C1, 1-2), 4Ch 1-4). 51, 1-2), o@2, 1-4), 701), 8, 1-3), 96, 2-7), 10d), 11(1), 14(1). Segment VIII: hair 0(1), 4(1), 9(3, 3-5), 14(1). Paddle: Uniformly light- ly pigmented, midrib darker; external margin serrated; hair 1-P slender. Male genital lobe extending to about 0.4 of paddle, female genital lobe to 0.35. LARVA (fig.48). Head: 0.84 mm. Siphon: 1.00 mm. Anal Saddle: 0.32 mm. Read- ily separated from guerrero by the presence of abdominal spicules on both dorsal and ventral surfaces and anal saddle with distinct large marginal spicules ventrad of hair 1-X. Head: Width about 1.06 of length. Pigmentation moderate except around mouthparts where it is deeper. Mental plate with a strong median tooth and usually 11(11,12) distinct teeth on either side, often a smaller detached tooth at base. Hair OC1).. 101). 301). 46..4-8), 5C1),6(2),. 7, 2-3), 8C1), 9(3,:3-5), OC, 1-3), t1a12,2- 17), 12(1, 1-2), 131, 1-2), 14(8, 4-9), 15(@2, 2-3). Antenna: Length about.0.37. of head. Pigmentation moderate; shaft with a few scattered spicules. All hairs except I, single; hair 1-A (2, 2-4) at about 0.63 from base. Thorax: Apices of long hairs atten- uate; barbs present on both long and stellate hairs. Prothorax: hair 0(24, 17-29), 1(6, 4-7), 2(1), 3(16, 11-19), 4(2, 1-2), 5(3, 2-3), 6(1), 7(3, 2-4), 8(28, 24-30), 9(28, 22- 28), 10(1), 11(2, 2-4), 12(1), 13(33, 22-35), 14(30, 24-38). Mesothorax: hair 1(11, — 6-14), 2(1), 3(1), 4(1), 51), 6(6, 4-7), 7(1), 8€6, 5-6), 9(7, 5-8), 10(1), 11@, 1-2), 12(1), 13(16, 10-17), 14(8, 8-13). Metathorax: hair 1(16, 12-25), 2(1), 365, 4-9), 4(17, 13-20), 5€15, 13-20), 6(1), 7(6, 3-8), 8(11, 10-14), 9(4, 3-6), 10(1), 111, 1-2), 12(1), 13(27, 20-33). Abdomen: Integument of segment I glabrous; other segments with spinelike spicules on dorsal and ventral surfaces. Stellate hairs with branches of unequal length. Segment I: hair 1(19, 12-20), 2(28, 17-28), 3(1), 4(6, 4-8), 5(16, 12- 19), 6(3, 2-3), 71, 1-2), 9(6, 5-9), 10(1),; 11(17, 13-21), 13(10, 8-10). Segment II: hair 0(1), 1(17, 11-20), 2(26, 17-26), 3(1), 4(6, 4-8), 5(16, 14-20), 6(3, 2-3), 7(9, 6- 11), 8(1, 1-3), 9(6, 4-9), 10(1), 11(1), 12(1), 13(9, 7-16). Segment III: hair 0(1), 1(18, 8-22), 2(24, 16-27), 3(1), 4(1), 5(17, 12-26), 6(2, 2-3), 709, 6-12), 8(1), 9(14, 10-18), 10(1), 111), 12(1), 13(6, 5-12), 14(1). Segment IV: hair 0(1), 1(14, 5-18), 2(23, 15-27), 301,.1-2), 401), 5(15,10-20), 6(@2, 2-3), 7€9, 6-13); 8C1); 9C16, 12-22), 10(1), 11(1), 12(1), 13(7, 4-12), 14(1). Segment V: hair 0(1, 1-2), 1(12, 5-15), 219, 15-25), 3(1), 4C1, 1-3), 5(15, 11-18), 6(2), 7(8, 6-11), 8(1), 9(16, 11-23), 10(1), 11(1), 12(1), 13(8, 4-10), 14(1). Segment VI: hair 0(1), 114, 6-15), 2(17, 13-24), 3(1), 4(1), 512, 10-19), 6(2, 2-3), 7(8, 6-11), 8(1), 9(14, 11-17), 10(1), 11Q1), 121), 13(8, 4-10), 14(1, 1-2). Segment VII: hair 0(1), 1(11, 6-16), 2(16, 11-20), 31), 4(1), 5(1 1 8-17), 6(9, 6-14), 7(1); 86, 3-7), 9013, 10-21), 1001),. 1101), 12), 13(,, 3- 14), 14(1, 1-3). Segment VIII: Comb scales in a single row, usually 14(12-20) in number; individual scale elongate, pointed apically and evenly fringed nearly to tip. Hair 0(1), 1(10, 8-20), 2(1), 3(5, 4-6), 4(1), 5(6, 4-8), 14(2, 1-4). Siphon: Index about 4.0-4.2; integument moderately pigmented, imbricate. Pecten extending to about 0.55, with 22(17-24) teeth; individual tooth pointed apically, with basal den- ticles. Hair 1(3, 2-3), 2(1), 3-5 not visible, 6(1), 7(1), 8(1), 9(1). Anal Segment: Sad- Berlin: Subgenus Howardina : 101 dle imbricate; moderately pigmented, with darker basal ring; caudal margin with spines of varying sizes dorsad and ventrad of hair 1-X. Gills subequal, about 2.0 of saddle length. Hair 1(3, 2-4), 2(4, 3-5), 3(2), 4a(4, 2-4), 4b(3, 3-4), 4c(4, 3-4), 4d@3, 2-4), 4e(2, 2-3). SYSTEMATICS. A. allotecnon is the most widespread and most common of the 3 allopatric species in the Allotecnon Group. Its known range extends from the southern end of the Sierra Madre Oriental in Mexico to central Panama. The only sig- nificant variation noted in the large series of specimens was the presence of a few sil- very scales on upper ppn in the 2 females from Mexico (MEX 14,16). Although this condition tends toward guerrero the 2 forms appear to be distinct in other adult fea- tures and the 3 larvae of allotecnon from Mexico are clearly different from those of guerrero and apparently indistinguishable from those of allotecnon populations in Central America. Only with additional material of all stages from Mexico will it be possible to determine the taxonomic status of this population and its relation to guerrero. The distribution of allotecnon appears to be discontinuous in Central America probably because of its apparent restriction to areas of cloud forest. Al- though not reported as yet from Guatemala and Honduras, it may occur there in suitable environments. BIONOMICS. The immature stages of allotecnon have been collected in epiphytic bromeliads at elevations above 5,000 feet, frequently in association with members of the Quadrivittatus Group. A few females have been taken in daytime biting-landing collections on human bait. ; DISTRIBUTION (fig.6). Cordilleras of Middle America in Mexico (Sierra Madre Oriental and Sierra Madre de Chiapas), possibly eastern Guatemala, El Salvador, probably Honduras, Nicaragua, Costa Rica and Panama (to Canal Zone) at elevations above 5000 ft. Material examined: 521 specimens; 43 6, 48 2, 136 pupae, 294 larvae; 114 individual rearings (24 pupal, 46 larval, 44 incomplete). COSTA RICA. Alajuela: Poasito, 14 Nov 1962, C.L. Hogue and W.A. Powder (CR 23), 5 Ip? (23-101,102,104,106,110), 1 pd (23-111), 2 p¥ (23- 112,113), 3 IP (23-103,107,108), 6 P, 6 p, Bone we | [UCLA] . Volcan Poas, 14 Apr 1938, H.W. Kumm, 1 d;1L, 41 [USNM] ; Summit, 14 Nov 1962, C.L. Hogue and W.A. Powder, 1 Ipd (20-110), 1 Ip? (20-109), 6 IP (20-102-106,108), 5 L (CR 20), 2 Ipd (21-103,106), 2 Ip? (21-109,110), 1 pd (21-112), 1 p? (21-101), 6 IP (21-102,104, 105,107,109,111), 25 L (CR 21); 9 July 1963, C.L. Hogue, 3 6, 2 9,4 P,4p,4L,41(CR 145) [UCLA]. Cartago: Cervantes, 9 Nov 1962, J.N. Belkin, C.L. Hogue and W.A. Powder, 1 L(CR 9), 3 IP (11-101-103), 2 Ipd (13-104,108), 3 pd (13-102,109,110), 3 p? (13-101-103), 3 IP (13-103, 105,106), 2 L (CR 13), 7 Ipd (14-102-105,107,111,112), 2 lp? (14-109,110), 1 pd (14-101), 1 IP (14-108), 2 L (CR 14); 9 Nov 1962, C.L. Hogue (CR 12), 1 lpg (12-106), 1 IP (12-105) [UCLA]. La Sierra, elev. 6300-8500 ft, 24 Nov 1962, C.L. Hogue and W.A. Powder, 1 IP (46-201), 1 Ipd (49-106), 4 Ip? (49-104,108-110), 4 IP (49-101-103,105), 1 p, 1 L, 11(CR 49), 1 pd (51-102), 1 p? (51-101), 2 IP (51-104,105), 10 L (CR 51), 2 Ip (52-101,109), 1 Ip? (52-104), 4 IP (52-102, 103,107,110), 21 L (CR 52), 1 IP (53-104), 1 9, 15 L(CR 54) [UCLA]. Orosi, 7 Dec 1962, CLL. Hogue and W.A. Powder (CR 83), 1 lp? (83-501), 1 L [UCLA]. Rio Macho, 7 Dec 1962, C.L. Hogue and W.A. Powder, 1 lp? (79-102), 1 pd (79-101), 2 Ipd (86-103,104), 1 pd (86-101), 1 L (CR 86) [UCLA]. Volcan Iraju, 15 Nov 1962, C.L. Hogue and W.A. Powder (CR 24), 2 pd (24- 101,108), 1 p? (24-106), 3 IP (24-102-104), 1 P, 1 1 [UCLA]. Heredia: Puerto Viejo, 8 Sept 1965, W.J. Hjort (CR 220), 1 Ipd (220-11) [UCLA]. San Jose: Villa Mills, 21 Nov 1962, C.L. Hogue and W.A. Powder (CR 39), 1 lp? (39-104), 3 p? (39-101-103), 3 L [UCLA]. EL SALVADOR. Laguna de las Ranas, Jan 1942, H.W. Kumm, 4 6, 1 , 2 L [USNM]. MEXICO. Chiapas: Sumidero Canyon, 17 Aug 1964, E. Fisher and D. Verity, 3 L (MEX 116), 1 ? (MEX 121) [UCLA]. Veracruz: Cordoba, Fortin de las Flores, 11 July 1964, E. Fisher, 1 ° (MEX 14) [UCLA]. Also from states of Hidalgo, Queretaro and San Luis Potosi (Vargas 1956:272). NICARAGUA. Jinotega, 3 July 1964, J.N. Belkin and A. Quinonez (NI-33C), 1 IP (33C-41) [UCLA]. 102 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 PANAMA. Bocas del Toro: Chiriquisito, Trail Caldera, 5 Nov 1965, P. Orjuela (PA 1004), 6? [UCLA]. La Zorra, elev. 4400-5400 ft, 6 Apr 1963, A. Quinonez, 1 L (PA 176); 7 Apr 1963, A. Quinonez, 5 L (PA 179); 8 Apr 1963, A. Quinonez, 4 L, 1 1(PA 180); 9 Apr 1963, A. Quinonez, 7 L (PA 183) [UCLA]. Chiriqui: Palo Santo, elev. 4600 ft, 15 June-1 Dec 1950, 1 2 (Trapido, Ga- lindo and Carpenter 1955:530). Boquete, 30 Jan 1914, J. Zetek, 1 @ [USNM]. El Hato del Vol- can, Chiriqui Lagoon, elev. 4500 ft, 12 Mar 1964, A. Quinonez (PA 638), 2 lp? (638-101,102), 1 IP (638-108), 1 L [UCLA]. Cocle: El Valle, elev. 2000 ft, 24 Aug 1963, A. Quinonez (PA 533), 1 p? (533-101), 2 L [UCLA]. Panama: Cerro Campana, 3 Aug 1963, A. Quinonez (PA 462), 1 lp? (462-108), 1 p? (462-101) [UCLA]. 25. Aedes (Howardina) guerrero Berlin, n.sp. Figs.6,9 49 TYPES: Holotype ? (MEX 166), 21 miles south of Chilpancingo, Guerrero, Mexico, 31 Aug 1964, E. Fisher [USNM]. Paratypes: 2 9 (MEX 166), same data as holotype [UCLA]. FEMALE. Wing: 3.06 mm. Proboscis: 2.34 mm. Forefemur: 2.16 mm. Abdomen: about 3.15 mm. In general similar to allotecnon, differing apparently only in the fol- lowing. Head: Erect scales on occiput whitish in middle, darker on sides. Palpus about 0.2 of proboscis, with white scales on apex and mesal side of segment 4. Thorax (fig. 9): Mesonotal light lines golden. Creamy scales on median scutellar lobes narrow dis- tally, broader basally. Paratergite with broad silvery scales. Scales on apn broad, sil- very; upper ppn with predominantly broad, silvery scales, creamy narrow ones re- stricted to upper border; lower ppn patch small, with broad silvery scales; psp usual- ly with a small silvery patch; other scale patches bright silvery. Legs: Foretarsus and midtarsus with short incomplete white rings on segments 1,2; hindtarsus with broad- er white rings on 1-3. Haltere: Stem pale, knob mostly creamy, with a few black bas- al scales. Abdomen: Basomedian tergal light scales white from segment III caudad, silvery on segments VI,VII. Sternites with broad basal V-shaped white scale patches, silvery on distal segments; apical white scales inconspicuous. MALE. Unknown. PUPA. Imperfectly known. Appears very similar to allotecnon but possibly differ- entiated by the hair 5-IV,V which is shorter, sorry extending beyond the apex of the following tergite. LARVA (fig.49). Head: 0.81 mm. Siphon: 0.7 mm. Anal Saddle: 0.3 mm. Read- ily separated from allotecnon by the presence of abdominal spicules only on the ven- tral surface and anal saddle without distinct marginal spicules ventrad of hair 1-X. Head: Width subequal to length. Pigmentation as in allotecnon. Mental plate with a strong median tooth and 13(12-14) distinct teeth on either side, often a smaller de- tached tooth at base. Hair 0(1), 1(1), 3(1), 465, 5-6), 5(1, 1-2), 6(2), 7(2), 8(1), 9Q, 2-4), 10(2, 2-3), 11(7, 5-8), 12(2, 1-2), 13(1), 14(5, 4-8), 15(4, 3-4). Antenna: Length about 0.33 of head; shaft pigmentation moderate, spicules not apparent. All hairs ex- cept 1 single; 1-A (2) at about 0.6 from base. Thorax: Integument glabrous as in allo- tecnon. Apices of long hairs attenuate; barbs present on both long and stellate hairs. Prothorax: hair 0(13, 11-15), 1(6, 3-7), 2(1), 3(13, 10-14), 4@, 2-3), 5C1, 1-2), 6(2), ~7(2), 8(21, 21-25), 9(17, 13-19), 10(1), 1102, 2-4), 12(1), 1315, 9-17), 1434, 30- 38). Mesothorax: hair 1(7, 4-7), 2(1), 3(1), 41), 5(1), 6(5, 5-6), 7(1), 8(7, 6-7), 9(8, 6-9), 10(1), 11(1), 12(1), 13(14, 10-16), 14(4, 3-7). Metathorax: hair 1(5, 4-6), 2(1), 3(5, 2-6), 4(9, 8-10), 5(9, 8-11), 6(1), 7(7, 6-8), 8, 6-11), 9(4), 10€1), 11€1), 12), Berlin: Subgenus Howardina 103 13(16, 11-17). Abdomen: Integument of segment I glabrous; other segments with spinelike spicules restricted to ventral surface; hairs 1,7,13-II-VII apparently not stel- late. Segment I: hair 1(7, 4-8), 2(11, 10-15), 3(1), 4(4, 4-5), 5(12, 10-13), 6(3, 3-4), 7(1), 9(6, 5-9), 10(1), 11€11, 10-15), 13(5, 4-7). Segment II: hair 0(1), 1(4, 4-7), 2(9, 8-12), 3(1), 4(4, 4-6), 5(11, 9-12), 6(3, 3-4), 7(5, 4-7), 8(1), 9(5, 5-7), 10(1), 11(1), 12(1), 13(4, 4-5). Segment III: hair 0(1), 1(4, 3-5), 2(11, 8-11), 3(1), 4(1), 5(11, 9-12), 6(2), 7(5, 4-6), 8(1), 9(7, 5-9), 10(1), 111), 121), 13(4, 3-5), 14(1). Segment IV: hair 0(1), 1(4, 3-5), 2(10, 7-11), 3(1), 41), 510, 10-13), 6(2), 75, 5- 7), 8(1), 9(8, 6-8), 10(1), 11(€1), 12(1), 13(4, 4-5), 14(1). Segment V: hair 0(1), 1(4, 3-4), 2(8, 8-10), 3(1), 4(1, 1-2), 5(10, 8-11), 6(2), 7(5, 5-8), 8(1), 9(7, 5-9), 10(1), 11(1), 121), 1305, 3-6), 14(1). Segment VI: hair 0(1), 1(3, 3-5), 2(8, 5-9), 31), 4(1), 5(9, 6-11), 6(2), 7(4, 3-5), 8(3, 3-4), 9(7, 6-9), 10(1), 11€1), 121), 13(5, 4-5), 14(1). Segment VII: hair 0(1), 1(3, 3-4), 2(8, 7-10), 3(1), 41), 5(7, 5-9), 6(3, 2-4), 7(1, 1-2), 8(5, 5-7), 9(5, 3-9), 10(1), 111), 12(1), 13, 3-5), 14(1). Segment VII: Comb scales in a single row, usually 14(12-16) in number; individual scale elongate, pointed apically, evenly fringed almost to tip. Hair 0(1), 1(9, 4-11), 2(1), 3(7, 6-9), 4(1), 5(6, 5-7), 14(1). Siphon: Index about 3.3-3.6. Integument imbricate, moder- ately to strongly pigmented, with darker basal ring. Pecten extending to 0.42, usual- ly with 18(17-22) teeth; individual tooth pointed apically, with basal denticles. Hair 1(3, 2-4), 2(1), 3-5 not visible, 6(1), 7(1), 8(2, 1-2), 9(1). Anal Segment: Saddle im- bricate, moderately to strongly pigmented, with darker basal ring; caudal margin with spicules present only dorsad of hair 1-X. Gills subequal, long, more than 2.0 of saddle. Hair 1(4, 3-5), 2(5, 5-7), 3(2), 4a(4, 3-5), 4b(3, 3-4), 4c(3, 3-4), 4d(3, 2-3), 4e(2, 1-2). SYSTEMATICS. Although closely resembling allotecnon, this form appears to be a distinct species restricted to the Sierra Madre del Sur in Mexico where no other member of the group has been collected. The association of the immature stages with the females is presumptive only as no adults were obtained from the rearings and were found in a different locality from the adults but is probably correct since the larva is obviously of the allotecnon type but with distinctive features as indicated in the diagnosis. The pupal stage is not described at this time as it is known only from 2 whole imperfect specimens from incomplete rearings. It also appears to be general- ly similar to allotecnon but possibly differentiated by the shorter hair 5-IV-V. BIONOMICS. The presumptive immature stages of guerrero have been collected in epiphytic bromeliads at elevations of about 7,200-7,800 feet (MEX 429,430,431, 434). The 3 females were taken in a biting-landing collection (MEX 166) at 0800 in a pine-oak woodland. DISTRIBUTION (fig.6). Sierra Madre del Sur in Mexico at elevations above 7000 ft. Material examined: 20 specimens; 3 9, 2 pupae, 15 larvae; 2 incomplete individual rearings. MEXICO. Guerrero: Chilpancingo, 21.0 mi S, 21 Aug 1964, E. Fisher, 3 ? (MEX 166, type se- ties) [USNM, UCLA]. Oaxaca: Ixtlan de Juarez, El Punto, elev. 7700 ft, 13 Aug 1966, D.A. Schroe- der (MEX 434), 1 IP (434-10); Vivero Rancho Teja, elev. 7200 ft, 12 Aug 1966, D.A. Schroeder, 1 IP (429-10), 7 L (MEX 429), 1 L (MEX 430), 5 L (MEX 431) [UCLA]. 26. Aedes (Howardina) guatemala Berlin, n.sp. Figs.6,9 TYPES: Holotype ? (GUA 138-13), Yepocapa, Chimaltenango, Guatemala, 20 July 1950, Mi- 104 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 guel Xinic and Pedro Garcia [USNM]. Paratypes: 4 9? (GUA 138-10,12,14,15), same data as holo- type [UCLA]. FEMALE. Wing: 3.2 mm. Proboscis: 2.37 mm. Forefemur: 2.01 mm. Abdomen: about 2.6 mm. Very similar to allotecnon, differing apparently only in the following. Head: Erect scales on occiput predominantly dark, median ones lighter. Thorax (fig. 9): Mesonotal light lines pale golden. Lateral scutellar lobe with a few narrow dark and pale golden scales. Upper ppn scales predominantly creamy, narrow, a few scales slightly broader at lower border. Legs: Foretarsus and midtarsus with a few white scales on segment 1; hindtarsus with short white rings on segments | and 2 only. Haltere: Stem pale, knob predominantly white, dark scales restricted to base. MALE, PUPA, and LARVA unknown. SYSTEMATICS. The 6 known females of guatemala are so different in the de- tails of ornamentation from other members of the Allotecnon Group that I do not hesitate in recognizing them as members of a distinct species even in the absence of immature stages. This species appears to be restricted to the Pacific slope of central Guatemala where no other species of Howardina has been recorded. _ BIONOMICS. The immature stages will probably be found in epiphytic bromeli- ads. The females were probably taken in a biting-landing collection on human bait. DISTRIBUTION (fig.6). Pacific slope of Guatemala. Material examined: 6 speci- mens; 6 2; no individual rearings or immature stages. GUATEMALA. Chimaltenango: Yepocapa, 20 July 1950, Miguel Xinic and Pedro Garcia (GUA 138, type series), 5 2 (138-10,12,13-15); 24 July 1950, V.C. and R. Feoli, 1 @ (GUA 139) [USNM, UCLA]. Eleanorae Group FEMALES. Head: Median light line on vertex usually with narrow scales (moder- ately broad in eleanorae). Proboscis distinctly longer than forefemur. Torus with a restricted mesal scale patch. Thorax: Mesonotal light lines usually creamy to golden (except in ecuadoriensis in which inner dorsocentral and lateral marginal lines are white); inner dorsocentral line complete; outer dorsocentral line usually shorter, not reaching anterior margin, subparallel to inner; lateral marginal line continuous; scales on anterior promontory creamy, golden or white. Antealar area above paratergite with golden scales. Paratergite with a moderate scale patch extending entire sclerite length. Upper ppn entirely covered with light scales; upper stp patch quadrate. Legs: Distal white spot on anterior surface of midfemur distinct; present or absent on hind- femur. Tibiae completely dark. Wing: Veins entirely dark scaled. Abdomen: Basome- dian tergal light markings absent or present on III-VII. MALES. Palpus with or without basomedian light scales on segments 4 and 5. MALE GENITALIA. Lobe of tergite IX prominent or indistinct, with 4-7 strong bristles. Sidepieces separated at base by a distance less than width of aedeagus, with- out any indication of lobes on tergomesal or sternomesal margins. Claspette repre- sented by a conical tubercle. Aedeagus shorter than or subequal to proctiger; apical emargination distinct or shallow, lateral lobes conspicuous or inconspicuous. PUPAE. Cephalothorax: Trumpet moderate, index 4.5-6.4, not expanded apical- ly; hairs 10,12-C varied in branching. Abdomen: Hair 5-IV-V moderate to long, some- times exceeding apex of second tergite following; hair 9-VII moderate, reaching or exceeding lateral angle of tergite. Paddle: Moderate; apex usually rounded. Berlin: Subgenus Howardina | 105 LARVAE. Head: Hairs 4-7-C varied in branching; 15-C moderate to long, always extending beyond base of mental plate. Thorax: Roundish in outline; spiculation dense, weak or absent, spicules hairlike, simple. Abdomen: Integument strongly or weakly uniformly spiculate. Hair 8-VII distinctly cephalodorsad of 9-VII. Siphon: In- tegument smooth or imbricate; acus distinct, separate. Anal Segment: Hair 3-X single or double; ventral brush with 5 pairs of hairs. DISCUSSION. The Eleanorae Group of 6 species is probably derived from the Quadrivittatus Group and seems to replace the Allotecnon Group south of Panama. The ornamentation of the midfemur and hindfemur is quite varied in this group, more or less bridging the characteristic development in the other 2 groups. The lar- vae are also quite varied and somewhat intermediate between the Quadrivittatus and Allotecnon Groups, although the ventral brush always has only 5 pairs of hairs as in the latter group. Members of this group have been misidentified in the past as quad- rivittatus. The Eleanorae Group appears to be entirely confined to the Andes where it has been collected from moderate elevations up to nearly 11,000 feet. The group is now known from western and possibly northcentral Venezuela, Colombia, Ecuador and Bolivia (fig.6). Although seemingly some of the species are found sympatrically, there are indications of strong ecological divergence in other instances (ecuadoriensis and brevivittatus for example). This group is apparently sympatric to a limited ex- tent with the Whitmorei Group in the north and with the Whitmorei and Aurivitta- tus Groups in the south. 27. Aedes (Howardina) eleanorae Berlin, n.sp. Figs.6,9,50,5 1 TYPES: Holotype ¢ (COB 10-21) with associated pupal and larval skins and genitalia slide, Buritica, Antioquia, Colombia, 5000 ft, 8 Jan 1965, E. Osorno-Mesa [USNM]. Allotype ° (COB 10- 20) with associated pupal and larval skins, same data as holotype [USNM]. Paratype ? (COB 10- 22) with associated pupal and larval skins, same data as holotype [UCLA]. This species is dedicated to my wife, Eleanor, for her interests in and contribution to the study of this group. Aedes (Howardina) quadrivittatus in part of Dyar (1928:229); Edwards (1932:155); Lane (1939: 100; 1953:713); Reyes (1955:74,89); Stone, Knight and Starcke (1959:174). FEMALE. Wing: 3.06 mm. Proboscis: 2.34 mm. Forefemur: 2.07 mm. Abdomen: 2.88 mm. Head: Scales on median stripe predominantly broad and white, a few ante- rior ones narrow; light scales on sides and venter white. Erect scales on occiput and sides creamy. Palpus about 0.16 of proboscis. Mesal light scale patch on torus creamy. Thorax (fig.9): All mesonotal light lines creamy, of uniform width throughout; outer dorsocentral line short, arising at anterior 0.25 and reaching scutellum; lateral mar- ginal line long, from side of anterior promontory to wingbase, appearing broad from scutal angle to wingbase owing to presence of creamy scales in antealar area above paratergite; scales on anterior promontory creamy. Median scutellar lobe with broad dark scales laterally and broad creamy scales in the middle from base to apex; lateral lobes with a few narrow creamy scales. Scales on paratergite moderately broad, white. Apn scales broad, white, a few anterior ones narrower, creamy; upper ppn predomi- nantly with narrow, curved scales, a few broader scales below; lower ppn patch dis- tinct and merging with ssp patch; other scale patches pure white, distributed as in the 106 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 group. Legs: Forefemur dark anteriorly, predominantly dark posteriorly except for a creamy longitudinal line; midfemur with narrow anteromesal creamy line to 0.67 terminating in a white spot, posteriorly proximal 0.5 creamy, rest dark; hindfemur creamy to 0.67 on both sides, rest dark scaled. Foretarsus with a few white scales on segment 1; midtarsus with short incomplete white ring on 1; hindtarsus with broader white rings on 1 and 3, second segment dark. Haltere: Stem pale, knob predominant- ly white, a few dark scales dorsally. Abdomen: Basomedian tergal light scales absent. Sternites with broad V-shaped white patches; apical light scales inconspicuous. MALE. Wing: 2.88 mm. Proboscis: 2.43 mm. Forefemur: 1.89 mm. Essentially similar to female in color. Palpus entirely dark. MALE GENITALIA (fig.50). As figured; diagnostic characters as in the key. Ninth tergite lobe not prominent, with 4,5 strong bristles. Sidepiece length about 3.0 of median width. Clasper about 0.5 of sidepiece, with 1,2 subapical hairs; basal 0.3 with minute spicules; spiniform long, about 0.5 of clasper. Aedeagus shorter than proc- tiger; dorsal sclerotization emarginate apically, producing lateral sclerotized lobes. PUPA (fig.50). Abdomen: 3.3 mm. Trumpet: 0.39 mm; index 4.4-4.5. Paddle: 0.67 mm. Readily separated from other members of the group (except Jeei) by the strongly developed hair 8-VII on dorsal surface; from Jeei by the branched hair 10-C. Chaetotaxy as given below, hairs moderately to darkly pigmented. Cephalothorax: Mesonotum lightly pigmented between trumpets, lateral areas and metanotum dark- er; trumpet moderately pigmented and sharply contrasting. Hair 1(1), 2(1), 3(1, 1-2), 4(3, 2-5), 5(2), 6(1, 1-2), 7(2), 81, 1-3), 9(1, 1-2), 10(6, 5-6), 11(1, 1-2), 12(4, 2-5). Abdomen: Tergite I moderately pigmented except for usual membranous areas; II- VII moderately pigmented, progressively lighter caudad. Segment I: hair 1(13-16 pri- mary branches), 2(1, 1-3), 3(1), 4(2, 2-4), 5(1, 1-2), 6(1), 71, 1-2), 9(1, 1-3), 11Q, 2-3, present in some). Segment II: hair 0(1), 1(6, 4-7), 2(¢1), 3(1), 4(2, 2-3), 5(2, 1- 2), 6(1), 7(2, 1-3), 9(1), 10 (present in a few). Segment III: hair 0(1), 1(2, 2-4), 2(2, 1-3), 31, 1-2), 4C1, 1-2), 5(1, 1-2), 6(1), 71, 1-2), 8(1, 1-3), 9(1), 10€1), 111), 14(1). Segment IV: hair O(1), 1(2, 2-3), 2(1, 1-2), 3(3, 2-3), 4(1, 1-2), 5(2), 6(1), 71, 1-2), 8(1, 1-2), 9C1), 101), 11€1), 14(1). Segment V: hair 0(1), 1(2), 201, 1-2), 3(2, 1-2), 4(2, 2-3), 5(2), 6(1), 7(2, 1-3), 8(@2, 1-3), 9(1), 10(1), 111), 141). Seg- ment VI: hair O(1), 1@, 1-2), 2(@2, 1-2), 3(1, 1-2), 4(2, 1-2), 5(1), 6(1), 71, 1-2), 8, F279. C1), FOLT), (1), P4C1) Segment Vi: hair OC), 1(1, 1-2), 201,142). 3,12), 4(1), 51, 1-3), 6(3, 1-3), 7(1), 8(4, 3-5), 9(4, 4-8), 10(1), 11(1), 14(1). Segment VIII: hair 0(1), 4(1), 9(11, 11-14), 14(1). Paddle: Lightly pigmented, midrib slightly darker; hair 1-P single. Male genital lobe extending to 0.5 of paddle, female genital lobe to 0.34. LARVA (fig.51). Head: 0.63 mm. Siphon: 0.28 mm. Anal Saddle: 0.27 mm. Diag- nostic characters as in the key. Readily separated from other members of the group by pecten teeth extending beyond hair 1-S. Head: Width subequal to length; pigmen- tation moderate except around mouthparts where it is deeper. Mental plate with a strong median tooth and usually 12 distinct teeth (11-12) on each side, often a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 4(12, 9-15), 5(8, 6-9), 6(5, 4-6), 7(17, 14-18), 8(12, 10-14), 9(14, 13-16), 10(2, 2-3), 11(34, 20-42), 12(10, 7-10), 13(1), 14€18, 18-22), 15(2, 2-3). Antenna: Length about 0.5 of head; shaft uniform- ly lightly pigmented and with few scattered spicules; hair 1(3, 2-3), others single. Thorax: Integument densely covered with hairlike simple or branched spicules except on intersegmental areas; long hairs barbed and with attenuate apices; stellate hairs with many weakly barbed branches. Prothorax: hair 0(30, 27-34), 1(5, 3-5), 2(1), 3(42, 35-42), 4(2, 1-2), 5(1), 6(1), 7(2), 8(30, 28-34), 9(22, 18-24), 10(1), 112, 2- Berlin: Subgenus Howardina 107 3), 12(1), 13(36, 31-36), 14(52, 48-62). Mesothorax: hair 1(27, 23-32), 2(1), 3(1), 4(1), 5(1), 6(4, 3-5), 7(1), 8(4, 4-6), 9(5, 5-7), 10(1), 1101, 1-2), 12(1), 13(22, 18- 23), 14(22, 18-26). Metathorax: hair 1(37, 34-40), 2(1), 3(23, 20-26), 4(22, 15-25), 5(21, 17-23), 6(1), 7(4, 4-5), 8(23, 23-35), 9(3, 3-4), 10(1), 111, 1-3), 12(1), 13(26, 24-30). Abdomen: Integument and stellate hairs as on thorax. Segment I: hair 1(28, 25-28), 2(28, 26-30), 3(1), 4(1), 5(24, 22-25), 6(3, 2-3), 7(1), 9(12, 10-15), 10(1), 11(24, 20-24), 13(30, 29-35). Segment II: hair 0(1), 1(30, 29-31), 2(30, 24-32), 3(1), 4(1), 5(29, 25-30), 6(3), 7(20, 19-29), 8(1), 9(17, 16-17), 10(1), 11(1), 12(1), 13(36, 30-40). Segment III: hair 0(1), 1(30, 28-32), 2(31, 26-34), 3(1), 4(1), 5(28, 25-30), 6(2), 7(22, 17-23), 8(1), 9(22, 21-25), 10(1), 11), 12(1), 13(30, 30-33), 14(1). Seg- ment IV: hair 0(1), 1(32, 27-32), 2(29, 26-35), 3(1), 4(1), 5(30, 24-32), 6(2), 7(22, 18-22), 8(1), 9(26, 22-27), 10(1), 11¢€1), 12(1), 13(28, 26-29), 14(1). Segment V: hair O(1), 1(28,28-31),:2(30, 28-33), 3(1), 4€1), 5(30, 23-31); 6(2), 7022, 16-25), 8(1), 9(26, 25-26), 10(1), 11(1), 12(1), 13(30, 29-33), 14(1). Segment VI: hair 0(1), 1(30, 30-32), 2(28, 28-33), 3(1), 41), 5(30, 27-30), 6(1), 7(20, 12-21), 8(1), 9(26, 22-26), 10(1), 11(1), 12(1), 13(26, 23-38), 14(1). Segment VII: hair 0(1), 1(30, 28- 33), 2(30, 24-30), 3(1), 4(1), 5(29, 24-29), 6(14), 7(1), 8(24, 20-25), 9(24, 18-24), 101, 1-2), 11€1), 12(1), 13(28, 24-28), 14(1). Segment VIII: Comb scales in a single row, usually 12(12-14) in number; individual scale long, pointed apically and fringed almost to tip. Hair 0(1), 1(33, 28-34), 2(1), 309, 8-9), 4(1), 5(32, 27-35), 14(1). Si phon: Index about 3.3; integument imbricate, moderately pigmented except for darker basal ring. Pecten teeth usually 21(19-23), extending beyond hair 1-S to about 0.7 of siphon; individual tooth with basal denticles. Hair 1(4, 3-5), 2(1), 3-5 not visible, 6(1), 7(1), 8(1, 1-2), 9(1). Anal Segment: Saddle moderately uniformly pigmented except for darker basal ring; integument with minute spicules; caudal mar- gin with spines of varying length. Gills subequal, moderately long, about 2.0 of sad- dle length, narrow. Hair 1(2, 2-3), 2(4, 3-4), 3(1), 4a(2), 4b(2), 4c(2), 4d(2), 4e(1). SYSTEMATICS. A. eleanorae is the most strongly differentiated member of the group as indicated in the diagnosis above and in the keys. At present it is known from only 2 localities in Colombia, Buritica in the northern part of the Cordillera Occidental, and Buena Vista in the upper part of the Orinoco basin on the eastern slope of the Cordillera Oriental. Although eleanorae may be broadly sympatric with osornoi and marinkellei, it has not been found associated with these species and ap- parently occurs at lower elevations. BIONOMICS. The breeding sites of eleanorae are in epiphytic bromeliads at ele- vations of about 5,000 feet (COB 10). The female from Buena Vista was probably taken in a biting-landing daytime collection. DISTRIBUTION (fig.6). Upper Magdalena and Orinoco drainages in Colombia. Material examined: 15 specimens; | 6, 3 9, 4 pupae, 7 larvae; 3 individual larval rear- ings). COLOMBIA. Antioquia: Buritica, elev. 5000 ft, 8 Jan 1965, E. Osorno-Mesa (COB 10, type se- ries), 1 Ipd (10-21), 2 Ip? (10-20,22); 15 Jan 1965, E. Osorno-Mesa, 1 p, 1 L, 31 [USNM, UCLA]. Meta: Villavicencio, Buena Vista, 4 June 1942, W.H.W. Komp, 1 9? [UCLA]. 28. Aedes (Howardina) marinkellei Berlin, n.sp. Figs.6,9,52,53 TYPES: Holotype ? with associated pupal and larval skins (COM 63-19), Suba, Bogota, Cun- dinamarca, Colombia, 8300 ft, 16 Apr 1966, C.J. Marinkelle [USNM]. Paratypes: 1 9 (63-15) with 108 - Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 associated pupal and larval skins, 7 larvae, same data as holotype [UCLA]. This species is dedi- cated to Dr. C.J. Marinkelle for his contributions to the study of neotropical mosquitoes. Aedes (Howardina) quadrivittatus in part of Dyar (1928:229); Edwards (1932:155); Lane (1939: 100; 1953:713); Reyes (1955:74,89); Stone, Knight and Starcke (1959:174). FEMALE. Wing: 3.85 mm. Proboscis: 3.06 mm. Forefemur: 2.19 mm. Abdomen: about 3.41 mm. Similar to eleanorae, differing chiefly in the following. Head: Light scales on median stripe and short orbital line narrow, creamy; light scales on sides and venter broad, creamy. Erect scales predominantly creamy, darker on sides. Pal- pus about 0.16 of proboscis; predominantly dark, apex and mesal surface of segment 4 white scaled. Mesal patch on torus creamy. Thorax (fig.9): All mesonotal light lines creamy; outer dorsocentral line complete, broader on anterior 0.5, merging with lat- eral marginal line at level of lateral prescutal area; lateral marginal line long, moder- ately broad, extending from area of humeral bristle to wingbase; scales on anterior promontory creamy. Light scales on median scutellar lobe broad, white from base to apex; lateral lobes with a few narrow dark scales. Scales on paratergite moderately broad, creamy. Apn scales predominantly broad, dingy white, a few anterior ones nar- rower; ppn with only an upper patch of predominantly narrow creamy scales, a few lower ones whitish, broader; upper stp patch larger than in eleanorae; other scale patches white, distributed as in eleanorae. Legs: Forefemur predominantly dark an- teriorly except for a few scattered whitish scales along whole length, posterior sur- face mostly creamy except for a narrow dark line from base to apex; midfemur pre- dominantly dark anteriorly except for a few basal creamy scales and a white spot at distal 0.67, posteriorly mostly creamy with scattered dark scales on distal 0.5; hind- femur creamy to 0.67 on both sides, rest dark scaled. Foretarsus and midtarsus with basal white scales on segments 1,2; hindtarsus with broader white rings on segments 1-3. Haltere: Stem whitish, knob predominantly white, a few basal scales dark. Ab- domen: Basomedian tergal light scales dingy white. Sternites with large basal patches of dingy white scales; median apical white scales distinct to VI, on segment VII few and indistinct. Cercus with a few apical white scales. MALE unknown. PUPA (fig.52). Abdomen: 4.6 mm. Trumpet: 0.51 mm; index about 4.5. Paddle: 0.86 mm. Chaetotaxy as given below, hairs moderately to darkly pigmented. Distin- guished from osornoi by metanotal hairs all single, from /eei and eleanorae by ven- trally placed hair 8-VII. Cephalothorax: Mesonotum moderately pigmented between trumpets, lateral areas and metanotum darker; trumpets strongly contrasting. Hair 11), 21), 3(€1), 4(3), 5(2), 6(1), 7(2, 2-3), 8(1), 9(1), 10(1), 111), 121). Abdo- men: Tergite I moderately pigmented except for usual membranous areas; rest of seg- ments moderate, pigmentation progressively lighter caudad. Segment I: hair 1(6-7 primary branches, each with many secondary ones), 2(1), 3(1), 4(@, 1-2), 5(2), 6(1), 71), 91), 111). Segment II: hair 0(1), 12, 1-2), 2€1), 3C1), 4C1), 51), 61), 701), 9(1).. Segment LI: hair 0(1), 1(1,.4-2), 261), 3C1), 4€1,: 1-2); 5G), 6€1); 71, 1-2), 8(1), 9(1), 10(1), 11(1), 14(1). Segment IV: hair 0(1), 1(1, 1-2), 2(1), 3(2), 4Q, 1-2), 5(2), 6(1), 71, 1-2), 8(1), 91), 10€1), 11€1), 141). Segment V: hair 0(1), 1(1, 1-2), 2(1), SU), 41), S€1); 601): FO), SUT), OCT), TOG). 1101), T4C1).. Segment VT: hair O(1), 1€1), 2(1), 31), 4€1), 51), 6(1), 701), 8C1), 91), 10€1), 11¢1), 14€1). Segment VII: hair 0(1), 1(1), 21), 3(1), 41), 5(1), 6(1), 7(1), 81), 9(@2), 10(1), 111), 14(1). Segment VIII: hair 0(1), 4(1), 9(5, 5-7), 14(1). Paddle: Uniformly lightly pigmented, midrib darker; external margin serrated; apex rounded; hair 1-P slender. Female geni- tal lobe extending to 0.31 of paddle. Berlin: Subgenus Howardina 109 LARVA (fig.53). Head: 0.93 mm. Siphon: 0.81 mm. Anal Saddle: 0.32 mm. Chae- totaxy as given below, hairs moderately to darkly pigmented. Readily separated from all other members of Eleanorae Group by hair 5-C always double. Head: Width sub- equal to length; pigmentation moderate except around mouthparts where it is deeper. Mental plate with a strong median tooth and 10,11 distinct teeth on either side, of- ten a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 4(6, 5-6), 5(2), 6(3, 3-4), 7(5, 4-6), 8(1), 9(2, 2-4), 10(2), 11(12, 8-12; stellate, branches of varied lengths), 12(1), 13(1), 14(6, 4-8), 15(2). Antenna: Length about 0.33 of head; pigmentation moderate; shaft with scattered spicules. All hairs except 1 single; hair 1-A (2, 2-3) at about 0.58 from base. Thorax: Integument sparsely covered with simple, hairlike spicules. Apices of long hairs attenuate; long and stellate hairs weakly barbed. Pro- thorax: hair 0(17, 13-22), 1(3), 21), 3(20, 19-22), 4(2, 1-2), 5(1), 6(1), 7(2), 8(15, 10-16), 9(11, 9-13), 10(1), 111, 1-3), 121), 13(17, 16-20), 14(34, 34-38). Mesotho- rax: hair 1(7, 6-8), 2(1), 3(1), 4(1), 5(1), 6(4, 3-5), 7(1), 8(6, 5-6), 9(6, 4-8), 10(1), 11(1), 12(1), 13(10, 8-11), 14(2, 1-3). Metathorax: hair 1(9, 6-9), 2(1), 3(3, 2-4), 4(9, 8-10), 5(10, 7-10), 6(1), 7(6, 5-7), 809, 8-10), 9(3), 10(1), 11€1),.12(1), 1312, 10-18). Abdomen: Integument and stellate hairs as in thorax. Segment I: hair 1(13, £1517), 2(6, 8-11), 31), 401), 310, 9-16), 663, 2-3), 71), 9: 2-3), 1061); 1133. 11-14), 13(6, 4-8). Segment II: hair 0(1), 1(16, 12-16), 2(9, 8-11), 3(1), 4(1), 5(10, 8-12), 6(3), 7(6, 5-6), 8(1), 9(6, 5-7), 10(1), 11(1), 12(1), 13(9, 8-10). Segment III: hair O(1), 1(15, 11-16), 210, 10-12), 3(1), 4(1), 5(13, 11-14), 6(2), 7(8, 7-10), 8(1), 9(11, 6-11), 101), 11€1), 12(1), 1311, 8-12), 14(1). Segment IV: hair 0(1), 1(15, F3-17); 2(12; 9-13), Stl), 401), 3012, 10213); 6); 7, 70), B01). Stl 1-9-1 9). 10(1), 11€1), 121), 13€11, 8-12), 14(1). Segment V: hair 0(1), 1(14, 12-16), 2(10, £0-13),73(1);-4¢1), 52, 9-13), 00), 70769); Bly; OCT 1-15), 1001), Litt 12(1), 13(10, 9-11), 14(1). Segment VI: hair 0(1), 1(14, 11-16), 2(10, 9-12), 3(1), 4(1), 5(11, 10-13), 6(2), 765, 5-8), 8(1), 9(12, 10-14), 10(1), 11(1), 121), 1309, 7- 9), 14(1). Segment VII: hair 0(1), 1(10, 9-15), 2(10, 10-13), 3(1), 4(1), 5(10, 8-11), 6(5, 4-6), 7(1), 8(5, 5-6), 9(8, 5-10), 10(1), 11(1), 12(1), 13(5, 4-6), 14(1). Segment VII. Comb scales in a single row, usually 14(14-18) in number; individual scale long, pointed apically, fringed nearly to tip. Hair 0(1), 1(7, 7-11), 2(1), 3(5, 3-6), 4(1), 56, 6-9), 14(1). Siphon: Index about 3.0-3.2; moderately pigmented, imbricate. Pecten extending to about 0.5, usually with 16(16-19) teeth; individual tooth with basal denticles. Hair 1(1, 1-2), 2(1), 3-5 not visible, 6(1), 7(1), 8(1), 9(1). Anal Segment: Saddle moderately pigmented, with darker basal ring; caudal margin with spines of varying sizes. Gills subequal, at least 3.0 of saddle length, fingerlike. Hair 1(1), 2(4, 3-4), 3(2, 2-3), 4a(2, 2-3), 4b(2), 4c(2, 1-2), 4d(2, 1-2), 4e(2). SYSTEMATICS. A. marinkellei is most similar to osornoi and leei but is readily separated from both by constant diagnostic features in all known stages. It is known at present only from Cundinamarca Province in Colombia at elevations of over 8,000 feet where it is sympatric with osornoi. As indicated under the latter, a single reared female from this area combines features of osornoi in adult ornamentation and of marinkellei in the immature stages, suggesting hybridization between the 2 forms. Although found in the same general locality, osornoi and marinkellei have not been collected together in the same bromeliad axils. It appears, therefore, that some eco- logical barrier usually separates the 2 species. BIONOMICS. Immature stages of marinkellei have been found in the axils of epi- phytic bromeliads at elevations above 8,000 feet. Judging by the adult collection, fe- males of this species are apparently at least occasionally attracted to humans. DISTRIBUTION (fig.6). Cordillera Oriental in Colombia. Material examined: 22 110 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 specimens; 11 9, 2 pupae, 9 larvae; 2 larval individual rearings. | COLOMBIA. Cundinamarca: Bogota, 1918, Fr. Apollinaire-Marie, 1 2; B.G. Amortegui, 1 [USNM]. Rio San Francisco, 21 May 1942, W.H.W. Komp, 7 ° [USNM, UCLA]. Suba, elev. 8300 ft, 16 Apr 1966, C.J. Marinkelle (COM 63, type series), 2 lp? (63-15,19), 7 L [USNM, UCLA]. 29. Aedes (Howardina) leei Berlin, n.sp. Figs.6,9,54,55 TYPES: Holotype 2 (COL 17), 17.5 miles S.E. of Purace, Huila, Colombia, 10,000 ft, 28 Oct 1964, V.H. Lee [USNM] . Paratypes: 32 ? (COL 17), same data as holotype [USNM, BM, UCLA]. This species is dedicated to Dr. V.H. Lee, in recognition of his contributions to the study of neo- tropical mosquitoes. Aedes (Howardina) quadrivittatus of Martinez and Prosen €1953:28,30; 1955:33-33; 1959:603, 605); Martinez, Carcavallo and Prosen (1960:25); Reyes (1955:74,89); Prosen, Carcavallo and Martinez (1964:105). : FEMALE. Wing: 5.04 mm. Proboscis: 3.15 mm. Forefemur: 2.88 mm. Abdomen: about 4.32 mm. Similar to eleanorae but larger and differing primarily in the follow- ing. Head: Light scales on median stripe and short orbital line narrow, pale golden; light scales on sides and venter creamy. Erect scales dull creamy. Palpus about 0.17 of proboscis; predominantly dark, apex and mesal surface of segment 4 creamy. Mesal patch on torus creamy. Thorax (fig.9): All mesonotal light lines golden; outer dorsocentral line incomplete, extending from anterior 0.25 to scutellum; lateral mar- ginal line long, extending from side of anterior promontory to wingbase; scales on anterior promontory golden. Light scales on median scutellar lobe narrow, golden; lateral lobes with a few narrow dark scales. Scales on paratergite broad, creamy. Apn scales predominantly broad, creamy, a few anterior ones narrower, pale golden; ppn with only an upper patch of predominantly narrow golden scales; a few lower ones broader; other patches dingy white, distributed as in eleanorae. Legs: Forefemur dark anteriorly except for basal creamy scales, posterior surface with a creamy line broad to 0.5, narrowed distally, rest dark; midfemur mostly dark anteriorly except for basal creamy scales and a dingy white spot at distal 0.67, posteriorly proximal 0.6 creamy, rest dark; hindfemur creamy to 0.67 on both sides, rest dark scaled. Foretarsus with a few white scales on segment 1; midtarsus with short incomplete white rings on 1,2; hindtarsus with broader white rings on segments 1-3. Haltere: Stem pale, knob most- ly dark, fringe scales whitish. Abdomen: Basomedian tergal light scales distinct; din- gy white on segments II-VI, white on VII. Sternites with basal creamy patch; VI,VII with subapical band of white scales; apical light scales inconspicuous. MALE. Wing: 5.04 mm. Proboscis: 3.69 mm. Forefemur: 2.7 mm. Essentially sim- ilar to female in color. Palpus predominantly dark, with basoventral white scales on segments 4 and 5. : MALE GENITALIA (fig.54). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with 5,6 strong setae. Sidepiece length about 3.0 of median width. Clasper about 0.62 of sidepiece, with 2,3 subapical hairs; spiniform short, about 0.25 of clasper. Aedeagus subquadrate, longer than proctiger; dorsal sclerotiza- tion emarginate, producing lateral apical sclerotized lobes. PUPA (fig.54). Abdomen: 4.12 mm. Trumpet: 0.63 mm; index about 5.3. Paddle: 0.99 mm. Diagnostic characters as in the key. Chaetotaxy as given below, hairs mod- erately to darkly pigmented. Readily distinguished from eleanorae and osornoi by all Berlin: Subgenus Howardina 111 metanotal hairs single; from marinkellei by dorsally placed hair 8-VII. Cephalotho- rax: Mesonotum moderately pigmented between trumpets, lateral areas and metano- tum darker; trumpets strongly contrasting. Hair 1(1), 2(1), 3(1), 4(2, 2-3), 5(2), 6(1), 7(1, 1-2), 8(1), 91), 101), 1101), 12(1). Abdomen: Tergite I strongly pigmented ex- cept for usual membranous areas; rest of segments with moderate to strong pigmen- tation, progressively lighter caudad. Segment I: hair 1(9-12 primary branches, each with many secondary ones), 2(1), 3(1), 4(1, 1-2), 5(1, 1-2), 6(1), 7(1), 9(1). Seoment IT: hair 0(1), 1(2, 1-3), 3€1, 1-2), 4(2), 5(1), 6(1), 7(1), 8(1), 9(1). Segment ITI: hair 00), 101), 201), 301); 404), 5), 6€1), 7045 1-2),80), 91); 10014 104), 140) Sex ment IV: hair 0(1), 11, 1-3), 2(1), 31, 1-2), 4(1), 5(2), 6(1), 71, 1-2), 8(1), 9€1), 10(1), 11(1), 141). Segment V: hair 0(1), 1(1), 2(1), 3(1), 4(1), 5(2), 6(1), 7(2), 8(1), 9(1), 10(1), 11(1), 14(1). Segment VI: hair 0(1), 1(2), 3(1), 4(1), 5(1), 6(1), 71), 8C1), 9C1, 1-2), 1001), 111), 14€1). Segment VII: hair 0(1),:1(1; 1-2), 2(1), 3(1), 4(1), S(1), 6(2, 1-2), 7(1), 8(1), 9(4, 2-4), 10(1), 11(1), 14(1). Segment VIII: hair O(1), 4(1), 9(7, 6-7), 14(1). Paddle: Uniformly moderately pigmented, midrib darker; external margin serrated; apex faintly emarginate; hair 1-P slender. Male geni- tal lobe extending to 0.44 of paddle, female genital lobe to 0.31. LARVA (fig.55). Head: 0.94 mm. Siphon: 1.05 mm. Anal Saddle: 0.28 mm. Diag- nostic characters as in the key. Chaetotaxy as given below; hairs moderately to dark- ly pigmented. Readily separated from marinkellei by hair 5-C single; from osornoi by hair 3-X double; from eleanorae by pecten teeth terminating before hair 1-S. Head: Width subequal to length; pigmentation strong except around mouthparts where it is deeper. Mental plate with a strong median tooth and usually 12(12-14) distinct teeth on either side, often a smaller detached tooth at base. Hair 0(1), 1(1), 3(1), 4(7, 4-7), SY, T-2). 6(3, 2-3), 7(4, 4-5), 8€1, 1-2), 9(3), 10(3, 2-3), 11(16, 10-16, stellate, branches of varying length), 12(1), 13(1), 14(15, 10-19), 15(], '-2); Antenna: Length about 0.38 of head; pigmentation strong; shaft with a few scattered spicules. All hairs except I, single; hair 1-A (2, 2-3) at about 0.64 from base. Thorax: Integument weak- ly spiculate, spicules hairlike. Apices of long hairs attenuate; barbs present on long and stellate hairs. Prothorax: hair 0(26, 26-44), 1(2, 1-2), 2(1), 3(44, 34-72), 4(1, 1- 2), 5(1), 6(1), 7(2), 8(24, 23-36), 9(24, 13-34), 10(1), 11(1, 1-3), 12(1), 13(34, 29- 51), 14(36, 36-50). Mesothorax: hair 1(20, 9-32), 2(1), 3(1), 4(1), 5(1), 6(5, 4-6), 7(1), 8(6, 3-8), 9(7, 6-8), 10(1), 11€1), 12(1), 13(18, 15-29), 14(13, 3-21). Metatho- rax: hair 1(26, 21-44), 2(1), 3(4, 3-5), 4(18, 16-30), 5(16, 14-20), 6(1), 7(5, 3-7), 8(16, 14-20), 9(3), 10(1), 11(1, 1-2), 12(1), 13(26, 20-46). Abdomen: Integument and stellate hairs as on thorax. Segment I: hair 1(22, 21-40), 2(27, 17-37), 3(1), 4(1), S(14, 10-26), 6(2, 2-3), 7(1), 9(1, 1-2), 10(1), 11(20, 17-36), 13(19, 14-24). Segment II: hair 0(1), 1(22, 19-45), 2(23, 17-46), 3(1), 4(1), 5(16, 11-26), 6(3, 2-3), 7(11, 9- 17), 8(1), 9(9, 8-20), 10(1), 111), 12(1), 13(18, 16-23). Segment III: hair 0(1), 1(22, 22-50), 2(22, 18-47), 3(1), 4(1), 5(15, 14-32), 6(1, 1-2), 7(11, 8-20), 8(1), 9(20, 16-34), 10(1), 11(1), 12(1), 13(25, 16-36), 14(1). Segment IV: hair 0(1), 1(24, 21- 49), 2(22, 20-47), 3(1), 4(1), 5(16, 13-31), 6(1, 1-2), 7(10, 9-18), 8(1), 9(22, 18-33), 10(1), 11€1), 12(1), 13(25, 18-40), 14(1). Segment V: hair 0(1), 1(20, 18-46), 2(22, 18-45), 3(1), 41), 5(18, 14-26), 6(1), 7(10, 8-19), 8(1), 9(18, 16-31), 10(1), 11(1), 12(1), 13(24, 17-41), 14(1). Segment VI: hair 0(1), 1(24, 22-40), 2(22, 20- 40), 3(1), 4(1), 5(16, 11-24), 6(1), 7(8, 8-16), 8(1), 9(20, 18-31), 10(1), 11(1), 12(1), 13(16, 12-16), 14(1). Segment VII: hair 0(1), 1(24, 18-27), 2(18, 16-27), 3(1), 4(1), 5(16, 12-19), 6(8, 8-11), 71), 8(10, 10-14), 9(17, 15-21), 10(1), 11(1), 12(1), 13(15, 13- 13), 14(1). Segment VIII: Comb scales in a single row, usually 14(12-15) in number; individual scale long, pointed apically, fringed nearly to tip. Hair 0(1), 1(24, 19-24), 112 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 2(1), 3(7, 6-8), 4(1), 5(19, 11-21), 14(1). Siphon: Index about 3.8-4.3; integument imbricate, strongly pigmented, with darker basal ring. Pecten extending to about 0.5, usually 23(22-24) teeth; individual tooth with basal denticles. Hair 1(3, 2-3), 2(1), 3-5 not visible, 6(1), 7(1), 8(1), 9(1). Anal Segment: Saddle strongly pigmented, with darker basal ring; caudal margin with spines of varying sizes. Gills subequai, fin- gerlike, about 2.0 of saddle length. Hair 1(3, 2-4), 2(4, 3-5), 3(2), 4a(2), 4b(2), 4c(2), 4d(2), 4e(2, 1-2). | SYSTEMATICS. A. leei appears to be closely related to osornoi and marinkellei. It is sharply differentiated ecologically from the other 2 species, occurring apparent- ly at considerably higher elevations in the paramo. All the stages are known from the type locality of Paramo de Purace in southern Colombia at elevations of 9600- 10,500 ft (COL 16,17,19,52,53,54). The only other definite record is a single female from Zongo near La Paz, Bolivia. It seems very probable that this species will even- tually be found in the intermediate area in Ecuador and Peru. BIONOMICS. The immature stages have been collected in epiphytic and terres- trial bromeliads in the Paramo de Purace where a number of females were also taken biting-landing and entering a car between 1715 and 1815 hrs. DISTRIBUTION (fig.6). Paramo in southern Colombia and Bolivia, above 9000 ft. Material examined: 188 specimens; 1 6, 34 9, 31 pupae, 122 larvae; 6 individual rearings (2 pupal, 4 incomplete). BOLIVIA. Cochabamba: Chapare, Yungas del Palmar, elev. 3200 m, 1 ?(Prosen, Carcavallo and Martinez 1964:104). La Paz: Zongo, 1 Apr 1946, W.H.W. Komp, 1 ? [USNM]. COLOMBIA. Cauca: Paramo de Purace, 9600-10,500 ft, 18 Feb 1965, V.H. Lee (COL 52), 1 p? (52-101), 4 IP (52-18,19,29,36), 8 P, 38 L; 19 Feb 1965, V.H. Lee, 7 L (COL 53), 5 P, 36 L(COL 54) [UCLA]. Huila: 10.0-17.5 mi S.E. of Purace, 9600-10,500 ft, 28 Oct 1964, V.H. Lee, 1 pd (16-100), 33 2 (COL 17, type series); 30 Oct 1964, V.H. Lee, 9 P, 25L,11(COL 19),3P, 11 L (COL 20) [USNM, BM, UCLA]. 30. Aedes (Howardina) osornoi Berlin, n.sp. Figs.6,9 56,57 TYPES: Holotype ¢ (COB 37-16) with associated pupal and larval skins and genitalia slide, “Chorro de Padilla,’ Rio San Francisco, Bogota, Cundinamarca, Colombia, elev. 8200 ft, 14 July 1965, J.A. Ochoa [USNM]. Allotype 2 (COB 37-10) with associated pupal and larval skins, same data as holotype [USNM]. Paratypes: 2 6 (COB 37-11,12), 2 2 (37-17,18), with associated pupal and larval skins, same data as holotype [BM, UCLA]. This species is dedicated to Dr. E. Osorno- Mesa, in recognition of his important contributions to the knowledge of neotropical mosquitoes. Aedes (Howardina) quadrivittatus in part of Dyar (1928:229); Edwards (1932:155); Lane (1939: 100; 1953:713); Reyes (1955:74,89); Stone, Knight and Starcke (1959:174); Cova Garcia, Su- til and Rausseo (1966a:58,283; 1966b:12,141,171,334). FEMALE. Wing: 3.8 mm. Proboscis: 3.24 mm. Forefemur: 2.34 mm. Abdomen: about 3.8 mm. Similar to eleanorae but generally larger and differing mainly in the following. Head: Light scales on median stripe and short orbital line narrow, pale golden; light scales on sides and venter creamy. Erect scales predominantly pale gold- en, a few on sides darker. Palpus about 0.15 of proboscis; predominantly dark, with white scales on apex and mesal surface of segment 4. Mesal scale patch on torus creamy. Thorax (fig.9): All mesonotal light lines golden; outer dorsocentral line joined anteriorly to entire lateral prescutal area in fossa, gradually narrowed caudad Berlin: Subgenus Howardina 113 to scutellum; lateral marginal line separate only from scutal angle to wingbase; scales on anterior promontory golden. Light scales on median scutellar lobe predominantly narrow, pale golden, a few basal ones broader; lateral lobe with a few narrow dark and creamy scales. Scales on paratergite golden, moderately broad. Apn scales pre- dominantly broad, pale golden, but a few anterior scales narrower; ppn only with an upper patch of narrow golden scales; psp scales narrow, pale golden; other patches pure white and distributed as in eleanorae. Legs: Forefemur predominantly dark an- teriorly except for a few basal creamy scales, posterior surface mostly dark except for creamy longitudinal line from base to apex; midfemur dark anteriorly except for a few basal creamy scales and a dingy white spot at about 0.5, posterior surface creamy in basal 0.5 and along a narrow distal line, rest dark; hindfemur creamy in basal half. Foretarsus with white scales on segments 1,2; midtarsus with short incom- plete white rings on 1,2, occasionally a few white scales on 3; hindtarsus with broad- er white rings on 1-3. Haltere: Stem pale, knob predominantly creamy, a few dorsal scales dark. Abdomen: Basomedian tergal light scales absent. Sternites with broad basal V-shaped silvery white patches; apical white scales inconspicuous. MALE. Wing: 4.5 mm. Proboscis: 3.24 mm. Forefemur: 2.16 mm. Essentially sim- ilar to female in color. Palpus entirely dark. MALE GENITALIA (fig.56). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with 5-7 strong setae. Sidepiece length 2.5-3.0 of median width. Clasper about 0.67 of sidepiece, with 2 subapical hairs; spiniform moderate, length about 0.33 of clasper. Aedeagus subquadrate, shorter than proctiger; dorsal sclerotization shallowly emarginate, lateral lobes inconspicuous. PUPA (fig.56). Abdomen: 4.29 mm. Trumpet: 0.61 mm; index 6.3-6.4. Paddle: 0.94 mm. Chaetotaxy as given below, hairs moderately to darkly pigmented. Distin- guishable from Jeei and marinkellei by hairs 10,12-C branched; from eleanorae by the ventrally placed hair 8-VII. Cephalothorax: Mesonotum lightly pigmented between trumpets; lateral areas and metanotum slightly darker; trumpets strongly contrasting. Hair 1(1), 2(1), 31), 4(3, 1-4), 5@, 1-3), 6(1, 1-2), 7(2, 1-3), 8(2, 1-3), 9(1), 103, 1-4), 11(1), 12(@, 1-4). Abdomen: Tergite I moderately pigmented except for usual membranous areas; tergites II,III also moderately pigmented, distal segments pro- gressively lighter caudad. Segment I: hair 1(12-21 primary branches, subequal in length and each with a few branches), 2(1, 1-3), 3(1), 4(2, 2-3), 5(2, 1-3), 6(1), 7(1), 9(1), 111, 1-2 only occasionally present). Segment II: hair 0(1), 1(9, 3-19), 2(1, 1- 2), 3(1), 4(2, 1-3), 5C1, 1-2), 6(1), 7(1, 1-2), 8(1, 1-2), 9(1). Segment III: hair 0(1), as 2); 2171-3), SCl, 1-2); ACL 1-2), S01 42), CET): FO; 3-2) Be. 22 Os. 101), 1 1(1), 14G). Segment PV: hair 01); 12, 1-2); 2¢1, 1-2); 3(3; 2-3), 402); 50, 2-3), 6(1), 7(2, 1-2), 8C1, 1-2), 9(1), 10(1), 11(1), 14(1). Segment V: hair 0(1), 1(1), 211-3); 3€1), 40, 1-2), 5@2; 1-2), 6(1), 70; 1-3), 8@, 1-2); 901), 1001), 1101), 14(1). Segment VI: hair 0(1), 1(1), 21, 1-2), 3(1), 4€1, 1-2), 5(1, 1-2), 6(1), 7d, 1- 2), 8(2, 1-3), 9(1), 10(1), 111), 14(1). Segment VII: hair 0(1), 1(1), 2(1), 3(1), 4(1), S(1), 6(2, 1-2), 7(1), 8@, 1-3), 9(3, 2-4), 10(1), 11€1), 14(1). Segment VIII: hair 0(1), 4(1, 1-3), 9(8, 5-9), 14(1, 1-2). Paddle: Uniformly lightly pigmented, midrib darker; external margin serrated and apex rounded; hair 1-P slender. Male genital lobe ex- tending to about 0.47 of paddle, female genital lobe to 0.3. LARVA (fig.57). Head: 0.99 mm. Siphon: 0.91 mm. Anal Saddle: 0.33 mm. Read- ily separated from other members of the group by hair 3-X single. Chaetotaxy as given below. Head: Width subequal to length; pigmentation moderate except around mouthparts where it is deeper. Mental plate with a strong median tooth and 11 dis- tinct teeth on either side, often a smaller detached tooth at base. Hair 0(1), 1(1), 114 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 3(1), 4(6, 4-8), 5(1), 6(3, 3-4), 7(4, 4-5), 8(1), 9(2, 1-2), 10(1), 11°16, 9-16), 12(1), 13(1), 14(12, 7-16), 15(1, 1-2). Antenna: Length about 0.33 of head; pigmentation light, shaft with a few scattered spicules. All hairs except 1 single; hair 1-A (2, 2-3) about 0.64 from base. Thorax: Integument glabrous. Apices of long hairs attenuate; stellate hairs with unequal, weakly barbed, spikelike branches. Prothorax: hair 0(28, 24-33), 1(2),. 2011-2), 3G1,.27:36),.4(01), SC), 601), 7(2), 826, 20-29), 9016, 15- 21), 10(1), 11(2, 2-3), 12(1), 13(30, 29-37), 14(41, 38-45). Mesothorax: hair 1(22, 18-25), 2(1), 3(1), 4(1), 51), 6(4, 3-4), 7(1), 805, 4-6), 9(5, 4-7), 10(1), 11€1, 1-2), 12(1), 13(18, 13-20), 14(13, 11-15). Metathorax: hair 1(25, 23-37), 2(1), 3(16, 12- 16), 4(3, 3-8), 5(16, 14-19), 6(1), 7(4, 3-4), 8(16, 15-16), 9(3, 3-4), 10(1), 11€1, 1-2), 12(1), 13(20, 18-23). Abdomen: Integument weakly spiculate, spicules simple, hair- like; stellate hairs as on thorax. Segment I: hair 1(18, 18-21), 2(17, 16-22), 3(1), 4(1), 5(16, 13-18), 6(3, 2-3), 7(1), 9(6, 46), 10(1), 11016, 15-18), 13(16, 15-18). Segment II: hair 0(1), 1(20, 17-22), 2(20, 16-20), 3(1), 4(1), 5(16, 14-18), 6(2, 2-3), 7(12, 10-12), 8(1), 9(6, 6-9), 10(1), 11(1), 121), 13(24, 21-24). Segment III: hair O(1), 1(18; 18-22), 2017, 17-20), 3(1), 4(1), 5(16, 13-20), 6(1), 7(11, 10-13), 81), 9(14, 11-14), 10(1), 11(1), 121), 13(23, 21-28), 14(1). Segment IV: hair 0(1), 1(21, 19-24), 2(20, 16-22), 3(1), 4(1), 5(17, 16-19), 61, 1-2), 7€11, 10-13), 8(1), 9C15, 14-18), 10(1), 11(1), 12(1), 13(22, 18-25), 14(1). Segment V: hair 0(1), 1(21, 19- 22), 2(20, 17-22), 3(1), 41), 5(17, 16-19), 6(2, 1-2), 7010, 10-14), 8(1), 917, 14- 20), 10(1), 11(1), 12(1), 13(22, 20-25), 14(1). Segment VI: hair 0(1), 1(21, 19-21), -2(22, 17-24), 3(1), 4(1), 5(16, 14-18), 6(1), 7(12, 10-14), 8(1), 9(16, 14-19), 10(1), 11(1), 12(1), 13(24, 22-26), 14(1). Segment VII: hair 0(1) 1(22, 20-24), 2@20, 18- 22), 3(1, 1-2), 41), 5(18, 17-21), 6(9, 9-11), 71), 8(17, 14-19), 9(16, 14-17), 10Q1), 11(1), 12(1), 13(24, 20-27), 14(1). Segment VIII: Comb scales in a single row, usual- ly 16(14-16) in number; individual scale long, pointed apically, fringed nearly to tip. Hair 0(1), 1(24, 21-26), 2(1), 3(6, 5-7), 4(1), 5(24, 18-24), 14C1, 1-2). Siphon: Index about 3.3-3.67; integument moderately pigmented, with darker basal ring. Pecten extending to about 0.42-0.5, usually 18(16-22) teeth; individual tooth with basal denticles. Hair 1(4, 3-4), 2(1), 3-5 not visible, 6(1), 7(1), 8@2, 1-4), 9(1). Anal Seg- ment: Saddle moderately pigmented, with a darker basal ring; caudal margin with spines of varying sizes. Gills subequal, more than 2.0 of saddle length. Hair 1(2), 2(4, 3-4), 3(1), 4a(2, 2-3), 4b(2), 4c(2), 4d(2, 1-2), 4e(1, 1-2). SYSTEMATICS. A. osornoi is readily separated from its apparent nearest rela- tives, marinkellei and leei, by constant diagnostic features in all known stages. Imma- ture stages and females are known only from elevations above 8,000 feet in Cundina- marca Province, Colombia, in the central part of the Cordillera Oriental. A female collected at 9,000 feet in the Sierra Nevada of the Cordillera de Merida in Venezuela [USNM] is undoubtedly osornoi and it seems very likely that the record of quadri- vittatus from Aragua in the Cordillera de la Costa (Cova Garcia, Sutil and Rausseo 1966) also pertains to this species. In the fair sample of material no striking variation is apparent except for 1 reared female from Cundinamarca Province (COB 37-13) where osornoi is sympatric with marinkellei. In this specimen a suggestion of possi- ble hybridization or introgression of these 2 species is indicated in the osornoi-like ornamentation of the adult and in the resemblance of the immature stages to marin- kellei. In this area of sympatry, the 2 species have not been collected together in the same bromeliad axils, suggesting an ecological barrier of some kind between them. BIONOMICS. The known breeding sites of osornoi are epiphytic bromeliads at elevations above 8,000 feet. As in the case of marinkellei, several females have been taken, probably in biting-landing collections, at human bait. Berlin: Subgenus Howardina 115 DISTRIBUTION (fig.6). Colombia (Cordillera Oriental) and Venezuela (Cordillera de Merida and probably Cordillera de la Costa) at elevations of 8000-9000 ft. Materi- al examined: 114 specimens; 11 6, 45 9, 22 pupae, 36 larvae; 22 individual rearings (20 larval, 2 incomplete). COLOMBIA. Cundinamarca: Bogota, B.G. Amortegui, 5 ? [USNM]. Puerto Lievano and Bogo- ta, 6 Aug 1935, B.G. Amortegui, 9 9 [USNM]. Rio San Francisco, 21 May 1942, W.H.W. Komp, 7 2 [USNM]; “Chorro de Padilla,” elev. 8000 ft, 9 Feb 1965, E. Ruiz, 1 lpd (11-10), 1 lp? (11-11), 3 1(COB 11), 12 2, 10 L(COB 650209); 25 Feb 1965, E. Ruiz, 21(COB 14); 21 June 1965, J.A. Ochoa (COB 34), 3 Ipd (34-18,93,94), 5 Ip? (34-14-17,19), 2 lp (34-95,98), 1 3, 1 9; 14 July 1965, J.A. Ochoa (COB 36), 2 Ipé (36-21,22), 1 Ip? (36-20), (COB 37, type series), 3 Ipd (37-11, 12,16), 3 lp? (37-10,17,18) [USNM, BM, UCLA]. VENEZUELA. Aragua: (Cova Garcia. Sutil and Rausseo 1966b:141). Merida: Sierra Nevada, elev. 9000 ft, Dec 1923, E.P. de Bellard, 1 9? [USNM]. 31. Aedes (Howardina) ecuadoriensis Berlin, n.sp. Figs.6,9,58 TYPES: Holotype ? (ECU 4), Leon River, Saraguro, Azuay, Ecuador, 5500 ft, adult collection, 22 Mar 1965, L.E. Pena G. [USNM]. Paratypes: 4 9 (ECU 4), same data as holotype [UCLA]. FEMALE. Wing: 2.88 mm. Proboscis: 2.25 mm. Forefemur: 1.62 mm. Abdomen: about 2.25 mm. Similar to eleanorae, with the following diagnostic differences. Head: Light scales on median and short orbital lines moderately broad, white; sides and venter with white scales. Erect scales dark. Palpus about 0.15 of proboscis; predomi- nantly dark, with white scales on apex and base of segment 4. Mesal patch on torus white. Thorax (fig.9): Mesonotal light lines white and golden; inner dorsocentral line complete, narrow, white; outer dorsocentral line apparently complete, narrow, pale golden, extending from anterior margin to scutellum, a few anterior scales whit- ish; lateral marginal line short, whitish, extending from scutal angle to wingbase; scales on anterior promontory white. Light scales on median scutellar lobe broad, white, narrow and brown on lateral lobes. Scales on paratergite broad and white. Apn scales moderately broad, white; ppn with large upper and small lower scale patch; former with narrow white scales above, broad below; latter with broad white scales; other scale patches broad, white scaled, distributed as in the group. Legs: Forefemur dark scaled anteriorly, posterior surface predominantly creamy except for narrow longitudinal dark line from base to apex; midfemur and hindfemur mostly dark ante- riorly except for a ventral white line to middle and a white spot at 0.67, posteriorly proximal 0.5 creamy, rest dark. Foretarsus with short incomplete white rings on seg- ments 1,2; midtarsus with short white rings on 1-3; hindtarsus with broader white rings on 1-4. Haltere: Stem pale, knob predominantly white, a few brown scales dor- sally. Abdomen: Basomedian tergal light scales dingy white on II-VII. Sternites with broad basal V-shaped patches of white scales; apical light scales distinct up to seg- ment V. MALE. Wing: 3.06 mm. Proboscis: 2.71 mm. Forefemur: 1.75 mm. Essentially similar to female in color. Palpus predominantly dark, basoventral parts of segments 4 and 5 white scaled. MALE GENITALIA (fig.58). Diagnostic characters as in the key. Ninth tergite lobe prominent, with 6,7 moderate to strong bristles, inner markedly flattened and almost leaflike. Sidepiece length about 3.5 of median width. Clasper about 0.5 of sidepiece, with 2,3 subapical hairs; minute spicules in basal 0.5; spiniform short, 0.25 116 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 of clasper. Aedeagus short; dorsal sclerotization deeply emarginate, producing dis- tinct lateral sclerotized lobes. PUPA, LARVA unknown. SYSTEMATICS. A. ecuadoriensis is readily separated from the apparently closely related brevivittatus by constant differences in mesonotal, pleural and tarsal mark- ings. It is known only from 2 widely spearated localities in northern and southern Ecuador in semi-arid areas (chaparral and high desert) at elevations between 5,000 and 6,000 feet, whereas brevivittatus is known at present only from intermediate lo- calities with a range of elevations from 4,200 to nearly 11,000 feet, apparently al- ways in humid forests. BIONOMICS. Most of the known material of ecuadoriensis consists of females taken in biting-landing collections between the hours of 0600 and 1500. The imma- ture stages will probably be found in bromeliads which undoubtedly contain water during the period of seasonal rainfall characteristic of the areas where the adults were collected. DISTRIBUTION (fig.6). Ecuador (chaparral and high desert areas) at elevations of 5000-6000 ft. Material examined: 10 specimens; 2 6, 8 9; no individual rearings or immature stages. ECUADOR. Azuay: Saraguro, Leon River, elev. 5500 ft, 22 Mar 1965, L.E. Pena G., 5 9? (ECU 4), type series [USNM, UCLA]. Jmbabura: (barra, Taguando River, elev. 5400-5800 ft, 9 June 1965, LE. PenaG., 2 6, 3 9 (ECU 25) [UCLA]. 32. Aedes (Howardina) brevivittatus Berlin, n.sp. Figs.6,9,58 TYPES: Holotype ¢, Zorrocucho, Azuay, Ecuador, 3500 m, date not specified, Levi-Castillo [USNM]. Allotype ¢ with associated genitalia slide, same data as holotype [USNM]. Paratypes: 8 2, same data as holotype [USNM, UCLA]. Aedes (Howardina) quadrivittatus in part of Dyar (1928:229); Edwards (1932:155); Lane (1939: 100; 1953:713); Levi-Castillo (1953:40); Stone, Knight and Starcke (1959:174). FEMALE. Wing: 5.04 mm. Proboscis: 3.27 mm. Forefemur: 2.79 mm. Abdomen: about 4.55 mm. Similar to eleanorae, differing in larger size and in the following. Head: Light scales of median and short orbital lines creamy; light scales on sides and venter whitish. Erect scales predominantly brown, a few middle ones paler. Palpus about 0.14 of proboscis; predominantly dark, with white scales on apex and mesal side of segment 4. Mesal patch on torus white. Thorax (fig.9): All mesonotal light lines creamy to pale golden; inner dorsocentral line slightly broadened on anterior 0.5, gradually narrowing caudad; outer dorsocentral line shorter, from anterior 0.25 to scutellum; lateral marginal line longer, from anterior promontory to wingbase; scales on anterior promontory golden. Light scales on median scutellar lobe narrow, white; lateral lobes with narrow, brown scales. Scales on paratergite broad, dingy white. Apn scales broad, dingy white, a few anterior ones narrower; ppn with only an upper patch of predominantly narrow, creamy scales, a few broader ones below; oth- er pleural patches with broad white scales, distributed as in the group. Legs: Forefe- mur dark brown anteriorly, posteriorly proximal 0.5 predominantly creamy extend- ing distally as a narrow creamy line, rest dark; midfemur mostly dark brown anteri- orly except for a few basal creamy scales and a white spot at 0.67, posteriorly prox- Berlin: Subgenus Howardina 117 imal 0.5 creamy, extending distally as a creamy line, rest dark; hindfemur creamy to 0.75 anteriorly, 0.67 posteriorly, rest dark scaled. Foretarsus with a few white scales on segments 1,2; midtarsus with short incomplete white rings on 1-3; hindtarsus with short white rings on 1-4. Haltere: Stem pale, knob predominantly dark, fringe scales whitish. Abdomen: Basal tergal light bands creamy to V, white on VI,VII. Sternites with broad basal patches, creamy basally and white on sides; apical creamy to white scales distinct up to segment V. MALE. Wing: 5.6 mm. Proboscis: 3.45 mm. Forefemur: 2.37 mm. Essentially sim- ilar to female in color. Palpus predominantly dark, basoventral parts of segments 4 and 5 white scaled. MALE GENITALIA (fig.58). As figured; diagnostic characters as in the key. Ninth tergite lobe prominent, with 6,7 strong bristles. Sidepiece length about 2.5 of median width. Clasper about 0.5 of sidepiece, with 1,2 subapical hairs; spiniform moderate, about 0.33 of clasper. Aedeagus subquadrate; dorsal sclerotization deeply emar- ginate, producing distinct lateral lobes. PUPA, LARVA unknown. SYSTEMATICS. A. brevivittatus is apparently closely related to ecuadoriensis with which it is broadly sympatric in Ecuador but from which it seems to be isolated by a strong ecological barrier. While ecuadoriensis is known only from semi-arid lo- calities at elevations of 5,000 to 6,000 feet, brevivittatus has been collected only in humid forests at elevations from 4,200 to nearly 11,000 feet, apparently only on the western slopes of the Andes. BIONOMICS. The immature stages of brevivittatus will undoubtedly be found in epiphytic and/or terrestrial bromeliads. The present collections consist mostly of fe- males taken biting-landing on humans between the hours of 1000 and 1800. DISTRIBUTION (fig.6). Humid forests on western slopes of Andes in Ecuador at elevations of 4200-11,000 ft. Material examined: 25 specimens; 8 6, 17 9; no indi- vidual rearings or immature stages. ECUADOR. Azuay: Cuenca, Caserio Tinajillas, elev. 10,000 ft, 18-21 Mar 1965, L.E. PenaG., 7 3, 7 2 (ECU 3) [UCLA]. Zorrocucho, elev. 3500 m, Levi-Castillo, 1 6,9 9, type series [USNM, UCLA]. Canar: (Levi-Castillo 1953:40). Pichincha: Santo Domingo, Tandapi, elev. 4400-5400 ft, 20-21 June 1965, L.E. Pena G., 1 9? (ECU 27) [UCLA]. Aurivittatus Group FEMALES. Head: Median light line on vertex with narrow scales. Proboscis dis- tinctly longer than forefemur. Torus with restricted mesal scale patch. Thorax: Meso- notal light lines usually creamy to golden; inner dorsocentral line complete; outer dorsocentral line either complete or not reaching anterior margin, subparallel to in- ner; lateral marginal line continuous; scales on anterior promontory pale golden. An- tealar area above paratergite with golden scales. Paratergite with a moderate scale patch extending entire sclerite length. Upper ppn entirely covered with light scales; upper stp patch quadrate. Legs: Distal white spot on anterior surface of midfemur distinct, absent on hindfemur. Tibiae completely dark. Wing: Remigium with white scales. Abdomen: Basomedian tergal light markings absent. MALES. Palpus with basomedian light scales on segments 4 and 5. MALE GENITALIA. Lobe of tergite [X distinct, with 5,6 strong bristles. Side- pieces separated at base by a distance less than width of aedeagus; without any indi- cation of lobes on tergomesal or sternomesal margins. Claspette represented by a 118 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 conical tubercle. Aedeagus shorter than proctiger; apex slightly een pro- ducing lateral lobes. PUPAE and LARVAE unknown. DISCUSSION. The Aurivittatus Group is recognized here for the nominate form, vanemdeni and martinezi on the basis of the presence of pale scales on the remigium. The immature stages are unknown for the group and I have not seen any males. The group is doubtfully distinct from the Eleanorae Group and may have to be merged with it if the immature stages prove to be similar. Members of this group have been occasionally misidentified as sexlineatus in the past. The group occupies the southernmost portion of the distribution of the section. A. aurivittatus and vanemdeni have been found only in Bolivia, at moderate eleva- tions in the Amazon basin; martinezi is known at moderate elevations in the Amazon basin in Bolivia and from northern Argentina at an elevation of about 5,000 feet and is one of the 2 members of the section reported from the La Plata drainage (fig.5). 33. Aedes (Howardina) aurivittatus Cerqueira Figs.5 ,9 1943.Aedes (Howardina) aurivittatus Cerqueira, 1943a:7-8. TYPE: Holotype ?, Pampagrande, Florida, Santa Cruz, Bolivia, Feb 1940 [IOC]. Aedes (Howardina) aurivittatus of Cerqueira (1943b:33); Del Ponte, Castro and Garcia (195 1:240); Lane (1953:711); Stone, Knight and Starcke (1959:173); in part of Martinez, Prosen and Car- cavallo (1964:53-58); Prosen, Carcavallo and Martinez (1964:103); Forattini (1965:461); Stone (1967:208). FEMALE. Wing: 3.24 mm. Proboscis: 2.16 mm. Forefemur: 1.89 mm. Abdomen: about 2.88 mm. Head: Light decumbent scales on head white. Erect scales extending to vertex, light brown on occiput. Palpus about 0.16 of proboscis; white scales re- stricted to base and apex of segment 4. Mesal scale patch on torus silvery white. Tho- rax (fig.9): Mesonotal light lines creamy to pale golden; inner dorsocentral line com- plete, extending from anterior margin to scutellum; outer dorsocentral line complete, merging with lateral marginal line at level of humeral bristles; lateral marginal line long, extending from humeral bristles to wingbase. Median scutellar lobe with broad dark scales laterally and moderately broad creamy scales from base to apex in mid- dle; lateral lobe with a few narrow creamy scales. Paratergite with moderately broad white scales. Scales on apn predominantly broad and white, a few anterior ones nar- rower; ppn with large upper and small lower patches, former predominantly with narrow scales except for lower broader ones, latter with broad white scales; psp with a few broad white scales restricted to posterior part of bristles; other patches pure white, distributed as in the group. Legs: Forefemur mostly dark except for a few white scales at base anteriorly and a creamy longitudinal line posteriorly; midfemur predominantly dark anteriorly except for a narrow mesal line from base terminating in a distinct silvery spot at 0.67, posteriorly proximal 0.5 creamy, rest dark; hindfe- mur mostly creamy to 0.67 on both sides, rest dark scaled. Foretarsus and midtarsus with a few white scales on segment 1; hindtarsus with moderately broad white rings on segments 1-3. Haltere: Stem pale, knob brown scaled basally, fringe scales whit- ish. Abdomen: Sternites with broad V-shaped basal white patches; median apical white scales few, present up to segment V. Berlin: Subgenus Howardina 119 MALE. Not seen; according to the description of Martinez, Prosen and Carcavallo (1964:53-58), essentially similar to female as described above. MALE GENITALIA. Diagnostic characters as in the key. Ninth tergite lobe prom- inent, with 6 strong bristles. Sidepiece length about 3.0 median width. Clasper about 0.5 of sidepiece, with 1 subapical hair; spiniform long, about 0.5 of clasper. Aedeagus shorter than proctiger; dorsal sclerotization slightly emarginate, producing lateral lobes. | PUPA, LARVA unknown. SYSTEMATICS. The 3 females of aurivittatus | have examined are readily sepa- rated from martinezi by the diagnostic characters mentioned above. This species ap- pears to be restricted to moderate elevations in the upper parts of the Amazon drain- age in Bolivia. The record of aurivittatus from La Caldera (Salta), Argentina, is prob- ably erroneous and based on the closely related martinezi which has been collected in the same locality. : BIONOMICS. The immature stages are unknown but should be found in the axils of epiphytic and/or terrestrial bromeliads, the only known breeding sites of the mem- bers of the Sexlineatus Section. Females of aurivittatus are evidently at least occa- sionally attracted to humans, presumably in the daytime. DISTRIBUTION (fig.5). Bolivia (upper Amazon drainage) at moderate elevations. Material examined: 3 specimens; 3 9; no individual rearings or immature stages. BOLIVIA. Cochabamba: Chapare, 27 Apr 1944, 1 2 [USNM]. Santa Cruz: Florida, Feb 1940, N.L. Cerqueira, 2 ? paratypes [USNM]. 34. Aedes (Howardina) martinezi Berlin, n.sp. Figs.5 ,9 TYPE: Holotype ? (ARG-1), La Caldera, Salta, Argentina, 27 Mar 1966, A. Martinez [USNM]. Paratypes: 30 2, same data as holotype [USNM, BM, UCLA]. This species is dedicated to Dr. A. Martinez for his contributions to the study of neotropical mosquitoes. Aedes (Howardina) aurivittatus in part of Martinez and Prosen (1955:25-27); Castro, Garcia and Bressanello (1959:550). FEMALE. Wing: 3.42 mm. Proboscis: 2.36 mm. Forefemur: 2.19 mm. Abdomen: 2.91 mm. Essentially similar to aurivittatus, differing apparently only in the follow- ing. Head: Light decumbent scales dull creamy. Erect scales on occiput largely brownish, a few darker. Thorax (fig.9): Mesonotal light lines pale golden; outer dor- socentral line shorter, arising at anterior 0.25; lateral marginal line long, from side of anterior promontory to wingbase. Lateral scutellar lobes with a few narrow dark scales. MALE, PUPA, LARVA unknown. SYSTEMATICS. A. martinezi has been confused in the past with aurivittatus from which it is clearly differentiated by the diagnostic characters mentioned above which are remarkably constant in the small type series. Apparently martinezi extends from moderate elevations in the Amazon basin in Bolivia to the upper part of the La Plata drainage in northern Argentina. BIONOMICS. The immature stages are unknown but will probably be found in the axils of bromeliads. The adults have been captured biting and landing on humans during the day at elevations of about 5,000 feet. 120 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 DISTRIBUTION (fig.5). Bolivia (upper Amazon drainage) and Argentina (upper La Plata drainage), at moderate to high elevations. Material examined: 32 specimens; 32 2, no individual rearings or immature stages. ARGENTINA. Salta: La Caldera, 3.0 mi before mountain pass Sta. Laura, elev. 5000 ft, 27 Mar 1966, A. Martinez, 31 9 (ARG-1, type series) [USNM, BM, UCLA]. BOLIVIA. Cochabamba: Chimore, 30 Apr 1944,1 9 [USNM]. 35. Aedes (Howardina) vanemdeni Martini Figs.5 ,9 1931.Aedes vanemdeni Martini, 1931:208-209. TYPE: Lectotype 6, Yungas de Coroico (La Paz), Bolivia, elev. 1000 m [BM; selection of Mattingly 1955:31]. Aedes (Howardina) vanemdeni of Lane (1939:101); Martinez (1950:39); De] Ponte, Castro and Garcia (1951:240); Lane (1953:716); Martinez and Prosen (1953:28,30; 1955:32); Castro, Gar- cia and Bressanello (1959:550); Stone, Knight and Starcke (1959: 174): Prosen, Carcavallo and Martinez (1964:105). FEMALE. Not seen. MALE. Essentially similar to aurivittatus, differing apparently only in the follow- ing (fig.9). Lateral marginal line short, not connected to outer dorsocentral line at level of humeral bristles and extending from scutal angle to wingbase. Forefemur and midfemur with a distinct white spot anteriorly at distal 0.67. Foretarsus and midtar- sus with white scales on segments 1-3; hindtarsus with white ring only on segment |; rest dark scaled. Haltere: Stem pale, knob entirely white scaled. MALE GENITALIA. Ninth tergite lobe distinct, with 5 strong bristles. Sidepiece length about 3.0 of median width. Clasper and spiniform as in aurivittatus. PUPA and LARVA unknown. SYSTEMATICS. A. vanemdeni is readily differentiated from other members of the Aurivittatus Group by the distribution of the tarsal white markings. The male genitalia appear to be similar to aurivittatus. A. vanemdeni is known only by the 6 (lectotype) and 2 of the original type series from moderate elevations in the Amazon basin in Bolivia. BIONOMICS. The unknown immature stages will probably be found in bromeliad axils. Females are presumably at least occasionally attracted to man. DISTRIBUTION. Bolivia (fig.5). Material examined: d lectotype examined by J.N. Belkin [BM]. Berlin: Subgenus Howardina 121 REFERENCES CITED Aitken, T.H.G. 1960. A survey of Trinidadian arthropods for natural viral infections (August 1953 to December 1958). Mosquito News 20:1-10. Anduze, P.J. 1941. Lista provisional de los zancudos hematofagos de Venezuela (Diptera:Culici- dae). Bol. Entomol. Venezolana 1:6-18. 1943. Estudios de entomologia medica en el estado Merida-Venezuela. Bol. Ento- mol. Venezolana 2:189-196. 1947. Contribucion al estudio de los vectores de la fiebre amarilla en Venezuela. Acad. Cien. Fis. Mat. Natur., Caracas, Bol. 10:331-373. Antunes, P.C.A. 1937a. Informe sobre una investigacion entomologica realizada en Colombia. Co- lombia, Univ. Nac., Bogota. Fac. Med., Rev. 6:3-29. 1937b. Informe sobre una investigacion entomologica realizada en Colombia. Co- lombia, Univ. Nac., Bogota. Fac. Med., Rev. 6:65-87. Arnett, R.H. 1949. Notes on the distribution, habits, and habitats of some Panama culicines (Diptera:Culicidae). N.Y. Entomol. Soc., J. 57:233-251. 1950. Notes on the distribution, habits, and habitats of some Panama culicines (Diptera:Culicidae). N.Y. Entomol. Soc., J. 58:99-115. Bates, M. 1944. Observations on the distribution of diurnal mosquitoes in a tropical forest. Ecology 25:159-170. 1945. Observations on climate and seasonal distribution of mosquitoes in Eastern Colombia. J. Anim. Ecol. 14:17-25. 1949. The natural history of mosquitoes. New York, MacMillan. 379 p. Belem Virus Lab. 1964. Annual Report. Belem, Para, Brazil. 133 p. Belkin, J.N. 1955. The Tripteroides caledonica complex of mosquitoes in Melanesia (Diptera: Culicidae). Pacific Sci. 9:221-246. 1962. The mosquitoes of the South Pacific. v.1. Berkeley, U. Calif. Press. 608 p. 1968. Mosquito Studies (Diptera, Culicidae). IX. The type specimens of New World mosquitoes in European museums. Amer. Entomol. Inst., Contrib. 3(4): 1-69. Belkin, J.N., C.L. Hogue, P. Galina T.H.G. Aitken, R.X. Schick and W.A. Powder 1965. Mosquito Studies (Diptera, Culicidae). II. Methods for the collection, rear- ing and preservation of mosquitoes. Amer. Entomol. Inst., Contrib. 1(2):19-78. Belkin, J.N., R.X. Schick, P. Galindo and T.H.G. Aitken 1965. Mosquito Studies (Diptera, Culicidae). I. A project for a systematic study of the mosquitoes of Middle America. Amer. Entomol. Inst., Contrib. 1(2):1-17. Belkin, J.N., R.X. Schick and S.J. Heinemann 1965. Mociuité Studies (Diptera, Culicidae). V. Mosquitoes originally destined from Middle America. Amer. Entomol. Inst., Contrib. 1(5):1-95. Blanchard, R. 1905. Les moustiques. Histoire naturelle et medicale. Paris, F.R. de Rudeval. 673 p. 122 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Blanton, F.S. and E.L. Peyton 1958. Notes and distribution of Aedes mosquitoes in Panama based on a three year light trap survey. Mosquito News 18:30-33. Bonne, C. and J. Bonne-Wepster 1925. Mosquitoes of Surinam. Roy. Colon. Inst., Amsterdam, Afd. Trop. Hyg. 13:1-558. Bonne-Wepster, J. and C. Bonne 1920. Diagnoses of new mosquitoes from Surinam, with a note on synonymy. In- secutor Inscitiae Mens. 7:165-180. Boshell-Manrique, J. and E. Osorno-Mesa 1944. Observations on the epidemiology of the jungle yellow fever in Santander and Boyaca, Colombia, September 1941 to April 1942. Amer. J. Hyg. 40:170- 181. Bugher, J.C., J. Boshell-Manrique, M. Roca-Garcia and E. Osorno-Mesa 1944. Epidemiology of jungle yellow fever in Eastern Colombia. Amer. J. Hyg. 39:16-51. Carpenter, S.J., P. Galindo and H. Trapido 1952. Forest mosquito studies in an endemic yellow fever area in Panama. Mos- quito News 12:156-164. Carpenter, S.J. and E.L. Peyton 1952. Mosquito studies in Panama Canal Zone during 1949 and 1950 (Diptera, Culicidae). Amer. Midland Natur. 48:673-682. Castro, M., M. Garcia and M.D. Bressanello 1959. Diptera Culicidae Culicinae. Primeras Jornadas Entomoepidemiol. Argent., Buenos Aires, p.547-562. Causey, O.R. and H.W. Kumm 1948. Dispersion of forest mosquitoes in Brazil. Preiiminary studies. Amer. J. Trop. Med. 28:469-480. Causey, O.R., H.W. Kumm and H.W. Laemmert 1950. Dispersion of forest mosquitoes in Brazil. Further studies. Amer. J. Trop. Med. 30:301-312. Causey, O.R. and G.V. Dos Santos 1949. Diurnal mosquitoes in an area of small residual forests in Brazil. Entomol. Soc. Amer., Ann. 42:471-482. Cerqueira, N.L. 1943a. Algumas especies novas da Bolivia, e referencia a tres especies de Haemago- gus (Diptera, Culicidae). Inst. Oswaldo Cruz, Rio de Janeiro, Mem. 39:1-14. 1943b. Lista dos mosquitoes da Bolivia (Diptera, Culicidae). Inst. Oswaldo Cruz, Rio de Janeiro, Mem. 39:15-36. Coquillett, D.W. 1902. Three new species of Culex. Can. Entomol. 34:292-293. 1905. New Culicidae from West Indies and Central America. Entomol. Soc. Wash., Proc. 7:182-186. 1906a. Five new Culicidae from the West Indies. Can. Entomol. 38: 60-62. 1906b. A classification of the mosquitoes of North and Middle America. U.S. Bur. Entomol., B. Tech. Ser. 11:1-31. 1910. The type species of the North American Genera of Diptera. U.S. Nat. Mus., Proc. 37:499-647. Cova Garcia, P., E. Sutil and J.A. Rausseo 1966a. Mosquitos (Culicinos) de Venezuela, v.1. Caracas, Min. Sanid. Asistencia Berlin: Subgenus Howardina 123 Soc. 410 p. 1966b. Mosquitos (Culicinos) de Venezuela, v.2._ Caracas, Min. Sanid. Asistencia Soc. 406 p. Davis, N.C. and R.C. Shannon 1931. Further attempts to transmit yellow fever with aS of South Ameri- ca. Amer. J. Hyg. 14:715-722. Del Ponte, E., M.P. Castro and M. Garcia 1951. Clava para las especies de Psorophora y Aedes de la Argentina y comorcas vecinas. Diagnosis de Aedes (O.) raymondi n.sp. (Diptera, Culicidae). Soc. Cien. Argent., An. 151:228-243. Diaz Najera, A. | 1966a. Aedes de Mexico. Descripcion de una nueva especie del subgenero Howarda- ina (Diptera, Culicidae). Rev. Invest. Salud. Publ. (Mex.) 24:331-344. 1966b. OMe tropicales de Mexico. Rev. Invest. Salud. Publ. (Mex.) 26:57-64. Dyar, H.G. 1905a. Remarks on genitalic genera in the Culicidae. Entomol. Soc. Wash., Proc. 7:42-49. 1905b. A synoptic table of North American mosquito larvae. N.Y. Entomol. Soc., J,,43:22-26. 1905c. Brief notes on mosquito larvae. N.Y. Entomol. Soc., J. 13:26-29. 1906. Illustrations of mosquito larvae. Entomol. Soc. Wash., Proc. 8:15-21. 1918. The male genitalia of Aedes as indicative of natural affinities. Insecutor Ins- citiae Mens. 6:71-86. 1920. The Classification of American Aedes. Insecutor Inscitiae Mens. 8:103-106. 1925. The mosquitoes of Panama. Insecutor Inscitiae Mens. 13:101-195. 1928. The mosquitoes of the Americas. Washington, Carnegie Inst. (Publication 387). 616 p. Dyar, H.G. and F. Knab 1906a. Notes on some American mosquitoes with description of new species. Biol. Soc. Wash., Proc. 19:159-172. 1906b. The larvae of Culicidae classified as independent organisms. N.Y. Entomol. Soc., J. 14:169-230. 1907. Description of some American mosquitoes. N.Y. Entomol. Soc., J. 15:9-13. 1913. Three new Neotropical mosquitoes. Insecutor Inscitiae Mens. 1:76-78. 1919. New species of tropical American mosquitoes. Insecutor Inscitiae Mens. 7: 1-9. | Dyar, H.G. and M. Nunez Tovar 1927. Notes on biting flies from Venezuela. Insecutor Inscitiae Mens. 14:152-154. Edwards, F.W. 1932. Diptera fam. Culicidae. Jn P. Wytsman, Genera Insectorum, Brussells, Des- met-Verteneuil, Fasc. 194. 258 p. Fauran, P. 1961. Catalogue annote des Culicides signales en Guyane Francaise. Inst. Pasteur Guyane Franc. Inini, Arch. Publication 465. 66 p. Floch, H. and E. Abonnenc 1942a. Especes de moustiques signalees pour la premiere fois en Guyane Francaise. Inst. Pasteur Guyane Ter. Inini, Publication 41. 6 p. 1942b. Catalogue et distribution geographique des moustiques de la Guyane Fran- caise actuellement connus (1). Inst. Pasteur Guyane Ter. Inini, Publication 43. 10 p. 124 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 1944. Culicides et Ixodides Guyanais. Moustiques signales pour la premiere fois et presence de Ornithodorus talaje en Guyane francaise. Inst. Pasteur Guyane ~ Ter. Inini, Publication 86. 6 p. 1945. Les Moustiques de la Guadeloupe (II). Les Genres Megarhinus, Aedes, Culex, Deinocerites, Mansonia et Wyeomyia. Inst. Pasteur Guyane Ter. Inini, Publica- tion 110. 48 p. 1947. Distribution des culicines des genres autres que la genre Culex, en Guyane Francaise. Inst. Pasteur Guyane Ter. Inini, Publication 148. 12 p. Floch, H. and P. Fauran 1954. Lutte antipaludique et lutte antiamarile en Guyane francaise. Inst. Pasteur Guyane Franc. Inini, Arch. Publication 322. 67 p. Forattini, O.P. 1965. Entomologia Medica. v.2. Sao Paulo, Univ. 506 p. Galindo, P. 1964. Ecology of Arthropod-borne viruses in Panama. Gorgas Mem. Lab., Annu. Progr. Rep. 130 p. : Galindo, P., S.J. Carpenter and H. Trapido 193k: Ecological observations on forest mosquitoes of an endemic area in Panama. Amer. J. Trop. Med. 31:98-137. 1955. A contribution to the ecology and biology of treehole bisects mosquitoes of Panama. Entomol. Soc. Amer., Ann. 48:158-164. Galindo, P. and H. Trapido 1957. Forest mosquitoes associated with sylvan yellow fever in Nicaragua. Amer. J. Trop. Med. Hyg. 6:145-152. Galindo, P., H. Trapido and S.J. Carpenter 1950. Observations on diurnal forest mosquitoes in relation to sylvan yellow fever in Panama. Amer. J. Trop. Med. 30:533-574. Gerry, B.I. 1932. Morphological studies of the female genitalia of Cuban mosquitoes. Ento- Soc. Amer., Ann. 25:31-64. Giles, G.M. 1902. A handbook of the Gnats or Mosquitoes. London, John Dale and Daniels- son. 530 p. Gowdey, C.C. 1926. Catalogus Insectorum Jamaicensis. Jamaica. Dept. Agri., Ent. Bull. 4:1-114. Grabham, M. : 1906. Notes on some mosquitoes from Newcastle, Jamaica. Can. Entomol. 38: 167-173. 1907. Notes on some mosquitoes from Jamaica, West Indies. Can. Entomol. 39: 25-26. Hill, R.B. and C.M. Hill 1948. The mosquitoes of Jamaica. Inst. Jamaica, B., Sci. Ser. 4. 60 p. Horsfall, W.R. 1955. Mosquitoes, their bionomics and relation to disease. N.Y., Ronald Press. 123, Howard, L.O., H.G. Dyar and F. Knab 1912. The mosquitoes of North and Central America and the West Indies. v.1. Washington, Carnegie Inst. (Publication 159). 520 p. 1917. The mosquitoes of North and Central America and the West Indies. v.4. Washington, Carnegie Inst. (Publication 159). p. 525-1064. Berlin: Subgenus Howardina 125 Johnson, C.W. 1919. A revised list of Diptera of Jamaica. Amer. Mus. Nat. Hist., Bull. 41:421- 449. Knight, K.L. and E.N. Marks 1952. An annotated checklist of the mosquitoes of the subgenus Finlaya, genus Aedes. U.S. Nat. Mus., Proc. 101:513-574. Komp, W.H.W. 1936a. An annotated list of the mosquitoes found in the vicinity of an endemic focus of yellow fever in the Republic of Colombia. Entomol. Soc. Wash., Proc. 38:57-70. 1936b. The male and larva of Aedes dominicii Rangel & Romero Sierra, and the male of Aedes pseudodominicii sp.nov., representatives of a new subgenus (So- peria) of the genus Aedes, from Colombia. Entomol. Soc. Wash., Proc. 38:71- The # 1956. Notes on mosquitoes from an area of endemic yellow fever in Colombia. Entomol. Soc. Wash., Proc. 58:37-42. Kumm, H.W. and W.H.W. Komp 1941. Aedes (Howardina) allotecnon, a new species of Aedes from Costa Rica, and description of the larva, adult and male terminalia of Aedes quadrivittatus (Coq). Entomol. Soc. Wash., Proc. 43:17-25. Kumm, H.W., W.H.W. Komp and H. Ruiz 1940. The mosquitoes of Costa Rica. Amer. J. Trop. Med. 20:385-422. Kumm, H.W. and O. Novis , 1938. Mosquito studies on the Ilha de Marajo, Para, Brazil. Amer. J. Hyg. 27:498- ee oe Kumm, H.W. and H. Zuniga 1942. Mosquitoes of El Salvador. Amer. J. Trop. Med. 22:399-415. van der Kuyp, E. 1954. Mosquitoes of Netherlands Antilles and their hygienic importance. Stud. Fauna Curacao Caribbean Islands 5:37-114. _ Lane, J: 1936a. Notas sobre investigacoes entomologicas em localidades onde houve febre amarella sylvestre em Sao Paulo. Arch. Hyg. Saude Pub., Sao Paulo 2:127-133. 1936b. Notas sobre culicideos de Matto Grosso. Mus. Paulista, Sao Paulo, Rev. 20:173-206. 1939. Catalogo dos mosquitos neotropicos. Sao Paulo, Clube Zool. Brasil. 218 p. 1949. Zoogeography of the Culicidae in the world. Mus. Paranaense, Curitiba, Rev. 7:247-263. 1951. Synonymy of Neotropical Culicidae. Entomol. Soc. Wash., Proc. 53:333- 336. 1953. Neotropical Culicidae, v.2. Sao Paulo, Univ. p.554-1112. Levi-Castillo, R. 1952a. Estudios epidemiologicos sobre la Fiebre Amarilla selvatica en las Provin- cias de Manabi y esmeraldas de la zona costera de la Republica del Ecuador. Rev. Kuba Med. Trop. 8(10,11 y 12):76-81. 1952b. Vorlaufige liste der stechmucken (Uranotaeniini, Toxorhynchitini, Culi- cini, Aedini, and Sabethini) aus Ecuador (Diptera, Culicidae). Z. Tropenmed. Parasitol. 3(4):552-559. : 1953. Lista provisional y distribucion de los mosquitos culicinos en el Ecuador. Rev. Ecuator. Entomol. Parasitol. 1(1):34-45. 126 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 Lima, A. de Costa 1930. Sobre os mosquitos que se craim em buracos de arvores. Inst. Oswaldo Cruz, Rio de Janeiro, Mem. 23:255-260. Lutz, A. 1904. Catalogo dos culicideos Brasileiros e Sul-Americanos. /n Bourroul, C. Mos- quitos do Brasil. 16 p. 1905. Novas especies do Mosquitos do Brasil. Imprensa Med., Sao Paulo 13:65-70. Martinez, A.F. 1950. Algunos Culicidae nuovos o poco conocidos para las entomofaunas de Ar- gentina, Bolivia y Paraguay. Mision Estud. Patol. Reg. Argent. 21:33-41. Martinez, A.F., R.U. Carcavallo and A.F. Prosen 1960. Una nueva especie de Culicidae de Bolivia. Univ. Buenos Aires, Fac. Cien. Med., Cat. Microbiol. Parasitol., Rev. 28:25-29. Martinez, A. and A.F. Prosen 1953. Nuevos Culicidos para las entomofaunas de Argentina, Bolivia y Eaaaey: Mision Estud. Patol. Reg. Argent. 24:27-32. 1955. Las especies de Aedes (Howardina) de Argentina y Bolivia (Diptera, Culici- dae). Mision Estud. Patol. Reg. Argent. 26:21-41. 1959. Nuevos culicidos para las entomofaunas de Argentina, Bolivia y Paraguay. Primeras Jornadas Entomoepidemiol. Argent., Buenos Aires, p.603-607. Martinez, A., A.F. Prosen and R.U. Carcavallo 1964. El alotipo macho de Aedes (Howardina) aurivittatus Cerquiera. Inst. Med. Reg., An. 6(1/2):53-58. Martini, E.C.W. 1931. Ueber einige sudamerikanische Culiciden. Rev. Entomol. 1:199-219. 1935. Los mosquitos de Mexico. Mex. Dep. Salubr., Bo!. Tec. (A) 1. 65 p. Matheson, R. 1950. Medical Entomology. ed.2. Ithaca, N.Y., Comstock. 612 p. Mattingly, P.F. 3 1955. Mosquitoes (Diptera: Culicidae) from the Tropical Institute at Hamburg. Roy. Entomol. Soc. Lond., Proc. (B) 24:27-33. Montchadsky, A.S. and I. Garcia Avila 1966. Las larvas de los mosquitos (Diptera, Culicidae) de Cuba, su biologia y de- terminacion. Poeyana (A) 28. 92 p. Peryassu, A.G. 1908. Os culicidos do Brasil. Rio de Janeiro, Inst. Manguinhos. 407 p. Peyton, E.L., P. Galindo and F.L. Blanton 1958. Pictorial keys to genera of Panama mosquitoes. Mosquito News 15:85-100. Picado, C. 1913. Les bromeliacees epiphytes considerees comme milieu biologique. B. Biol. France Belg. (7) 47:215-360. Porter, J.R. 1967. A checklist of the mosquitoes of the Greater Antilles and the Bahama and Virgin Islands. Mosquito News 27:35-41. Prosen, A.F., R.U. Carcavallo and A. Martinez 1964. Culicidae de Bolivia. Inst. Med. Reg., An. 6(1/2):59-124. Reyes, P. 1955. Lista de mosquitos de Colombia, S.A. (Diptera, Culicidae). Soc. Biol. Bogo- ta, An. 7(2):46-94. Berlin: Subgenus Howardina 127 Root, F.M. 1927. Notes on the mosquito fauna of the Republic of Haiti. Amer. J. Hyg. 7: 463-469. Senevet, G. 1936. Les moustiques de la Martinique. Inst. Pasteur Algerie, Arch. 14:123-134. 1938. Les moustiques de la Guadeloupe. Inst. Pasteur Algerie, Arch. 16:176-190. Senevet, G. and L. Quievreux 1941. Les moustiques de la Martinique (2e memoire). Inst. Pasteur Algerie, Arch. 19:248-264. Shannon, R.C. 1931la. The environment and behavior of some Brazilian mosquitoes. Entomol. Soc. Wash., Proc. 33:1-27. 1931b. On the classification of Brazilian Culicidae with special reference to those capable of harboring the yellow fever virus. Entomol. Soc. Wash., Proc. 33: 125-164. Soper, F.L., H. Penna, E. Cordoso, J. Serafim, M. Frobisher and J. Pinheiroz 1933. Yellow fever without Aedes aegypti. Study of a rural epidemic in the Valle do Chanaan, Espirito Santo, Brazil, 1932. Amer. J. Hyg. 18:555-587. Spielman, A. and A.E. Weyer 1965. Description of Aedes (Howardina) albonotatus (Coquillett), a common do- mestic mosquito from the Bahamas. Mosquito News 25 :339-343. Stone, A. 1963. A Synoptic Catalog of the Mosquitoes of the World (Diptera, Culicidae). Supp. II. Entomol. Soc. Wash., Proc. 65:117-140. 1967. A Synoptic Catalog of the Mosquitoes of the World (Diptera, Culicidae). Supp. III. Entomol. Soc. Wash., Proc. 69:197-224. Stone, A. and K.L. Knight 1956. Type specimens of mosquitoes in the United States National Museum: II, The genus Aedes (Diptera, Culicidae). Wash. Acad. Sci., J. 14:213-228. Stone, A., K.L. Knight and M. Starcke 1959. A Synoptic Catalog of the Mosquitoes of the World (Diptera, Culicidae). Wash., Entomol. Soc. Amer. (Thomas Say Found. Publication 6). 358 p. Suarez, O.M. and P. Cova Garcia 1961. Estudio morphologio de Aedes (H.) ioliota Dyar and Knab, 1913. Rev. Bra- sil. Entomol. 10:17-24. Surcouf, J. and R. Gonzalez-Rincones 1911. Essai sur les Dipteres vulnerants du Venezuela. Primiere Partie. Dipteres Nematoceres vulnerants. Paris, Maloine. 320 p. 1912. Dipteres piqueurs et suceurs de sang actuellement connus, de la Republique de Venezuela. Arch. Parasitol. Paris 15:248-314. Theobald, F.V. 1901. A monograph of the Culicidae or mosquitoes. v.1. London, Brit. Mus. (Natur. Hist.). 424 p. 1903a. A monograph of the Culicidae or mosquitoes. v.3. London, Brit. Mus. (Natur. Hist.). 359 p. 1903b. Notes on the genus “‘Stegomyia”’ (Theobald), and its distribution. J. Trop. Med. 6:237-239. 1905. Diptera, fam. Culicidae. In P. Wytsman, Genera Insectorum, Brussells, Des- met- Vertemeuil. Fasc. 26. 50 p. (128 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 1907. A monograph of the Culicidae or a v.4._ London, Brit. Mus. (Natur. Hist.). 639 p. 1910. A monograph of the Culicidae or mosquitoes. v.5. London, Brit. Mus. (Natur. Hist.). 646 p. Theobald, F.V. and M. Grabham 1905. The mosquitoes or Culicidae of Jamaica. Kingston, Inst. Jamaica. 40 p. Thompson, G.A. | 1947. A list of the mosquitoes of Jamaica, British West Indies. Mosquito News 7: 78-80. 1956. Aedes stenei, a new species of mosquito from Jamaica, B.W.I., with rede- scription of a closely related species (Diptera, Culicidae). Mosquito News 16: 196-200. Trapido, H. and P. Galindo | 1957. Mosquitoes associated with sylvan yellow fever near Almirante, Panama. Amer. J. Trop. Med. Hyg. 6:114-144. Trapido, H., P. Galindo and S.J. Carpenter 1955. A survey of forest mosquitoes in relation to sylvan yellow fever in Panama Isthmian area. Amer. J. Trop. Med. Hyg. 4:525 “942. Urich, F.W. 1913. Mosquitoes of Trinidad. Agr. Soc. Trinidad Tobago, Proc. 13:525-530 (Pap. 574). Vargas, L. 1956. Especies y distribucion de mosquitos mexicanos no Anofelinos (Insecta, Diptera). Inst. Salub. Enferm. Trop., Rev. 16:19-36. Whitman, L. 1951. The Arthropod vectors of yellow fever. In G.K. Strode (ed.). Yellow Fever. New York, Mc Graw Hill. 710 p. Whitman, L. and P.C.A. Antunes 1937. Studies on the capacity of various Brazilian mosquitoes, representing the genera Psorophora, Aedes, Mansonia and Culex to transmit yellow fever. Amer. J. Trop. Med. 17:803-823. nARWN Berlin: Subgenus Howardina 129 FIGURES Distribution of the groups of Howardina Distribution of the species of the Fulvithorax Section Distribution of the species of the Walkeri Section Distribution of Aedes (H.) sexlineatus Distribution of the species of the Quadrivittatus, Whitmorei and Aurivittatus Groups Distribution of the species of the Allotecnon and Eleanorae Groups Biting-landing cycle of Howardina species near Porus, Jamaica Mesonotal scale pattern of Howardina species Mesonotal scale pattern of Howardina species Aedes (H.) fulvithorax; pleuron, male genitalia and pupa Aedes (H.) fulvithorax ; larva . Aedes (H.) septemstriatus; male genitalia and pupa . Aedes (H.) septemstriatus; larva . Aedes (H.) arborealis; male genitalia and pupa . Aedes (H.) arborealis; \arva . Aedes (H.) walkeri; adult morphology Aedes (H.) walkeri; pleuron, male genitalia and pupa . Aedes (H.) walkeri; larva . Aedes (H.) aurites; male genitalia and pupa . Aedes (H.) aurites; larva . Aedes (H.) grabhami; male genitalia and pupa . Aedes (H.) grabhami; larva . Aedes (H.) inaequalis; male genitalia and pupa . Aedes (H.) inaequalis; larva . Aedes (H.) stenei; male genitalia and pupa > . Aedes (H.) stenei; larva . Aedes (H.) ioliota; male genitalia and pupa . Aedes (H.) ioliota; larva . Aedes (H.) cozumelensis; male genitalia and pupa . Aedes (H.) cozumelensis; larva . Aedes (H.) albonotatus; male genitalia and pupa . Aedes (H.) bahamensis; male genitalia and pupa . Aedes (H.) bahamensis; larva . Aedes (H.) busckii; male genitalia and pupa . Aedes (H.) busckii; larva . Aedes (H.) quadrivittatus; pleuron, male genitalia and pupa . Aedes (H.) quadrivittatus; larva . Aedes (H.) lorraineae; \arva . Aedes (H.) brevis; male genitalia and pupa . Aedes (H.) brevis; larva . Aedes (H.) spinosus; male genitalia and pupa . Aedes (H.) spinosus; larva . Aedes (H.) sexlineatus; male genitalia and pupa . Aedes (H.) sexlineatus; larva . Aedes (H.) whitmorei; male genitalia and pupa . Aedes (H.) whitmorei; larva . Aedes (H.) allotecnon; male genitalia and pupa 130 Contrib. Amer. Ent. Inst., vol. 4, no. 2, 1969 48. Aedes (H.) allotecnon; \arva 49. Aedes (H.) guerrero; larva 50. Aedes (H.) eleanorae; male genitalia and pupa 51. Aedes (H.) eleanorae; larva 52. Aedes (H.) marinkellei; pupa — 53. Aedes (H.) marinkellei; larva 54. Aedes (H.) leei; male genitalia and pupa 55. Aedes (H.) leei; larva 56. Aedes (H.) osornoi; male genitalia and pupa 57. Aedes (H.) osornoi; larva 58. Aedes (H.) aureolineatus, pseudodominicii, ecuadoriensis and brevivittatus; male genitalia NOILOSS SNLVANITXSS SNLDJJIAUND NOILDJSS XVYOHLIA Ns La 4OWJIUM P pra y “pe o : ATS, 9D410UDa|a | ga XD4 a | &< ly ¢ ty —, ‘ : ;———*__ snp aulxas on os ie UOUDAIOI|D ‘. $0 SK o, UHAIN, % ESN. - : y > Se eat. ee Ps ? 4 ak aS 5 ee aN 7 SPSS Ssr— $ npOHALp onb RSE Saree NSS. nN Ne, he fi NE eS. nyosng——ti aeuer NS ES FE TLN, [Ogg coe SAN of SSreAY a : [1a4]DM 3 snjo }ouoq|D —— — ; ; D401)! : : , * NOILOSS IMSN WWM ets =. , ay Lu 9 3 bas he “es SS } i Pop ae “~ vs 4 ? Ys ree T ‘ “do ~ i gy ees “ es Z : 4 £ : Sy = 2 N | : Ri ,, ’ . ts “ig = VNIQUYVMOH SATIN ALALVLS 00s 0001 SYALIWOTIA VOIUANY WIGGIN = e Ps SXDIOUPAIN # te) ie % ! 2) i bd A PP." r Win 3 = Sy A k . TVG OY t tf 3 Dac) Sete <=> snjpeulloounD “Se, a Sa TI —— ool os 0 00s "GWALAWOTIN vortanW STC 110})0M a q2 iN snip4ouogio : SNLVLONOS IV ten ae , D{O}] O! SATIW ALNLVLS 00s 0001 SYaLAIWOTIA VOINHNV WIdd (em eau os SNLVSNITIXSS — i a SATIW ALALVLIS 90s Gol 0s 6 ocol 00s SHALANOTTA VOIUHINV HIGCIN os co ‘4 \ | ) 1 c s § oS ey x VAN 4 -w a - mx SNJO}JIALApONb 9D9UID140} lOJOWLIUM ah ISYOWLIHM 5) ne — IZoullIOW PFs ae 4 sn IND Petes. 7 ID} HAUND™ Petes. FP JUBPWAUDA “sx ty enV LIISA: | 7 : i ey = 2-S961-1N3-A90100Z-¥19N SHTIW ALALYLS 00S SYA LAWOTIA WV HI OO os 0 0010s 0 JX G 2 j CdIW ae hes ‘s SNLDIJIAIAGIG SISUsIOpDNDA -199| 2 tox 19|J94uUUDW 10UJOSO apsoudala U0UD94Oj|D DJOWe}onb NONOSLONNV | o1osionb 2S SSUNIGHYMOHS) Hy Qs. jy © Y ee .: a, ae CLA! \ \ \ \ \ \ Toua}s ("H) ‘Vv pue STjenboeul (“H) Yo We ay PIedeis (iH) y fo ee tee g SiInoy UI JWT], , ie dil ie \- 4 ‘B 7 a ao f Le Pd fy \ , / 4 \ \ at / / \ a \ Vv / Py = PN -.co \ \ ra ®, 7 < \ eg ‘a / \ i” / \ ] \ / { / ' / \ ! | / \ / \ ! ; 1 \ / 1 7® | a 1 @ 1 - \ / \ / \ / \ ee A oe: \/ r) Z 614 ° e* e eo? °° e °° ® VOIVNVE “SNuYOd UVAN SHIONdS VNIGUVMOH AO ATIOAO ONIGNV'I-DNILIG 0090 00SO OZ O€ OS 09 OZ 08 OTT Ocl yySneo seojyrnbsow jo JaquinNn HOWARDINA acrostichal Fig.8 inner dorsocentral outer dorsocentral lateral marginal lateral prescutellar Selle ek ie ne ate oe ioliota, cozumelensis albonotatus,bahamensis, lateral marginal median prescutellar antealar supraalar busckii quadrivittatus lorraineae HOWARDINA ‘@ ® °° = 5 sa Q » & 9° ne a a 2) = o ” @ ro) = > 2 B oO z = 8 = = os ~~ @D oo ; w ~ £ —_ o ® > ~~ = oO mo @ =< @ _> =< a Co = poe 2 yas , a x< co ro] > OE S — = Qo =} 3 Bo 5 = s . f: oD g ‘bb 5 Fl Ss 8 3 a 8 8 3 Ww ” = = — = & << 2 re = 2 o 2 fo% an = wo oO ® ) — g : : @ @ =. a Qa. r) inezi mart ittatus agurivt iensis ecuador HOWARDINA =e if \\( Wy, sey [ | oN fulvithorax _TR 591-121 Monos Island Trinidad HOWARDINA 6 apes mm if mi fulvithorax 5 4 TR 680-102 MG \K Verdant Vale Arima, St. George Trinidad ~HOWARDINA - cy septemstriatus HOWARDINA septemstriatus PA 301-107 Almirante Bocas del Toro Panama HOWARDINA ee, arborealis =e. >) HOWARDINA Aloesoebanja Upper Surinam Surinam Bis, pa arborealis 11 HOWARDINA — —— pops CLeoiitkhiadaL COSSSSSS SSS 2 2222neee2elZ0eZeee 24 SAAR PPLE LE SELES SESS SESESE SSS \y — Ry = “as postgenial plate 4 i 2 ee 1 fa , < pate IX tergite spermatheca VIII sternite D> Co ue See walkeri ~ 4'o EO iE, a o 2 | IGG 6 HOWARDINA walker} JA 379-10 Mt. Diablo amaica HOWARDINA i 10 7 4 3 . "Gi : 4 ) Vill booon abMRR: ‘At i iy = ; : L A), ie » 2 R/ y Wy Wy yy | : ths A ‘ Nii Man) Mn \ ai Lp | y hy iV yyy ] Ny hy Lani walkeri JA 243-25 sd, i) Porus Z| Hf \\N Manchester I 4 AN Jamaica 5 {| HOWARDINA | vie Fig.12 « x \ WwW S WW 1.0 r JA 682-101 aur t es Newcastle St. Andrew Jamaica HOWARDINA . 3 Wy * 0 4 sp 9 8 2 13 BRM gua css rN Bs 50 1 Proximal Distal Att 7 PT Ae ‘ M : ‘ 14 « VIII : \ JA 682 Newcastle St. Andrew Jamaica aurites HOWARDINA JA 632-103 Porus Manchester Jamaica HOWARDINA grabhami eu TUS WS WY 2 pass " Nae JA 290-11 Stony Hill St. Andrew : , St An inaequalis amaica HOWARDINA Temple Hall St. Andrew Jamaica inaequalis HOWARDINA stenei Hope Farm Porus, Manchester HOWARDINA = Iii x 4 4 : ee \Q H\\ Ih ; tf / \ Hh. \ NaN Z \ I \\ \ a xf stene| \V4jI \ A , y \y & ¢ i, ae Kanne St. Andrew \ TR 940-111 * Trinidad Arima HOWARDINA ioliota HOWARDINA TR 938-129 Arima St. Andrew Trinidad HOWARDINA Island of Cozumel Yucatan Mexico cozumelensis HOWARDINA cozumelensis HOWARDINA \\ Fig. S| re San Francisco Mts Dominican Republic albonotatus HOWARDINA bahamensis HOWARDINA BAH 1 Hunter Grand Bahama Bahama Islands bahamensis HOWARDINA Fig. 34 Aedeagus Guadeloupe population DOM 41-10 “s ; sak busckil St Pat Dominica HOWARDINA busckii 0 HOWARDINA fate Fig. 36 HOWARDINA mee 3) Fy A. | fl, 35 Density of horni-infested suckers as function of sucker stand density Aare POWIet. TUG pr ae a ue Do Moa bs ge eice es ue ale ie eee ae pitdy ares. of Nord (1962) and Fowler (1) 9G) oo ses bce, Wilke (sos euvin suty depron Com Be 46 Agrilus horni study area near Beechwood, Michigan. ... . «+ ce. + aoe) 46 Ratio of observed to expected numbers of horni egg-bearing suckers per height class plotted. as.atunction of sucker Weight ows ace scenes eevee! 51 Ratio of observed to expected numbers of horni egg-bearing suckers per diameter class plotted as a function of sucker diameter. .......... 51 Ratio of total eggs to expected number of horni egg-bearing suckers per height class plotted as a function of sucker height. ............. 52 Ratio of total eggs to expected number of horni egg-bearing suckers per diameter class plotted as a function of sucker diameter. .......... 52 Number of suckers and cumulative number bearing Agrilus horni eggs per hypothetical random Sample. Of LO0OE iss ew acd ecces cs ircemese cae be 04 Aedeagus, Agrilus granulatus liragus (dorsal side). ...........-. 56 Ovipositor of Agrilus anxius in extended and retracted positions...... 57 Probable internal position of A. anxius aedeagus during copulation. .... D8 oii & Ie rtiat OFeans Ol FEDYORUCTION O'S a eee kw ee es 58 ORCI EO ee te aan dae aloes ab a: dnibeaitin gh RU 59 IC et A VR ree oe a tac acins eoclns cdesie anova asa tmanas 59 A)! SRS tte IROL On OL HOCEARUB ee ee eh ae ee eee 61 Se, Cf eT WEMETAIE ge kt ee Ma eee ek a ee, bei we cae 63 OUI GOVT IICre LORCING Blam ela pa, Wo ay oe ne we ee ee lade 63 Miele Geee TOT ODBOTVIN Aer US OTR Cae ake ee lee ee wh eG 66 Antennal fluttering by an Agrilus anxius CS incopulo............. 67 Antennal stroking movements by an Agrilus granulatus liragus “ in copulo . 68 Striate areas of scape and front (Agrilus vittaticollis %)........... 69 SSE EO OE eee CLUE PEED a eK aeRO mes ao MCR A EO Re oe 70 Internal surface prosternal lobe in Agrilus horni. ........4.4+.-+.2--. 71 Audiospectrograph of stridulation by Agrilus granulatus liragus %..... 72 Phyletic tree for part of the Agrilus anxius group. ........2454-+ 26s 75 Penis Sper OF ee OPPO kV elie el pa SR ORC a ate ee 79 Elytra of Agrilus ater, A. guerini, and A. quadriguttatus niveiventris ... 79 Mayr’s model for evolution of host specific sibling species. ........ 91 INTRODUCTION Objectives Beginning in the summer of 1964 we endeavored to study the natural history and taxonomy of four sympatric Agrilus Megerle’ (Coleoptera: Buprestidae). These Agrilus were of particular interest because of their morphological sim- ilarities. Three of them, the bronze birch borer (Agrilus anxius Gory), the bronze poplar? borer (A. granulatus liragus Barter and Brown) and the aspen root girdler (A. horni Kerremans) had only been distinguished rather recently. The fourth species, ‘ee pensus Horn (= betulae Fisher) is distinct from the oth- ers in coloration, but very similar otherwise. The credibility of contentions that these taxa are very closely allied will be deliberated here in the light of: 1) A taxonomic study of these and related Agrilus, which included working both with specimens and literature. 2) An analysis of their larval habits, their ecological relationships to their hosts, and their population-limiting factors. Much of what is known about these aspects is taken from the literature and reinterpreted in the light of more recent findings, and pertinent writings on other Agrilus are also discussed. 3) Direct and indirect observations on the behavior of the adults in the field and laboratory. 4) A preliminary study of interspecific mating and a theoretical analysis of phylogeny, zoogeography, and speciation (or subspeciation). Historic Background THE BRONZE BIRCH BORER. A. anxius is probably the most notorious Ne- arctic species in the genus, and at one time was thought to be the cause of birch dieback. It was first recognized as a pest of shade trees and ornamental birches in the late 1800’s and was called ‘‘the bronze birch borer’’ by Chittendon (1898). Chittendon says that ‘‘one correspondent has expressed the belief that if radical measures are not adopted, the loss of every birch in the city of Buffalo in the near future is imminent. This insect has already destroyed the common white birch [Betula papyrifera| and... many of the cut-leaf and European white bir- ches. It even attacks trees planted but a year before. ’’ Most of the reports of damage came from the northeastern U. S., perhaps because more birches had been planted there as shade trees. Nevertheless, ‘The majority of authors credit Curtis (1825) with originating the name Agrilus. How- ever, Megerle (1823) published a price list in which the heading ‘‘Agrilus v. M. Bupres- tis)’’ appears. This simple listing credits Megerle with authorship, and Curtis (1825) merely designated one of Megerle’s included species, Buprestis viridis L., as the geno- type. a The term poplar will here be used to refer to species in all sections of the genus Populus. 3 ; Described as Agrilus liragus Barter and Brown (1949); status changed by Carlson (1969). 2 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 the problem was not confined to that region. Chittendon (1900), for example, notes that in 1899 ‘‘Prof. F. C. Newcombe of the University of Michigan, Ann Arbor, Mich., wrote that what was without doubt this species, and which he designated as the white birch borer, had been in that locality for two or three years and had killed half the white birches in the city of Ann Arbor. ’”’ Swaine(1918) first reported anxius killing trees in forests. This was the start of a devastating and puzzling epidemic of birch dieback which primarily affected Betula papyrifera, although some mortality of Betula lutea (= alleghani- ensis) was also ascribed to this affliction. By 1951, 67 percent of the birches in Maine had been killed, and 15 percent of those remaining were in a decadent state due to the effects of birch dieback and bronze birch borer attack (Nash, Duda, and Gray, 1951). Much research concerning the bronze birch borer and dieback in the 1940’s and 1950’s was directed at determining whether or not the beetle was causing the death of the trees. This question was hardly a new one then. Chittendon (1898) says ‘‘as in the case of injury ascribed to the two-lined chestnut borer [Agrilus bilineatus Weber] there is still a certain question of doubt as to whether or not this birch borer is really the primary cause of the death of the trees. Our correspondent is of the opinion that the injury in Buffalo is due primarily to the attack of this borer, since it has been observed attacking vigorous trees. ’’ The work of Anderson (1944), Nash et. al. (1951), and Barter (1957) shows, how- ever, that the insect played.a secondary role, usually killing those trees al- ready afflicted with birch dieback. Whether or not birch dieback was the pre- disposing factor in earlier ravages of the bronze birch borer, when it allegedly killed mostly shade trees, is dependent upon what birch dieback really is. It is evident that the shade trees in many cities were in a weakened condition in the late 1800’s, perhaps because of droughtiness. THE BRONZE POPLAR BORER. The habits of A. granulatus liragus adults were first observed and recorded by Lintner (1883) before anything was pub- lished about the habits of A. anxius, and of course well in advance of the estab- lishment of the fact that the poplar feeder and birch feeder were not one and the same. Chittendon (1898) says, ‘‘a very singular thing in connection with the occurrence of this borer in birch is that in spite of frequent search, extending over a period of two years, our correspondent has been unable to find this in- sect attacking any other tree than birch--a remarkable condition of affairs when we consider the numbers of observations by careful observers of its occurrence on poplar and willow. ”’ Smith (1949a) discovered that the A. anxius (sensu Fisher, 1928) reared from Betula spp. had 22 chromosomes, while those reared from Populus spp. had only 20. Subsequently, Barter and Brown (1949) named the poplar feeding form A. liragus. Although A. granulatus liragus has habits nearly identical to those of anxius (host specificity excepted) it has never been accorded any great significance as a pest. Like anxius the bronze poplar borer attacks and kills only trees in weakened condition (Barter, 1965). because poplars are used less frequently as ornamentals than birches, g. liragus is not as impor- tant an ornamental pest as anxius. Furthermore, no large scale maladies on the order of birch dieback occur in forests of native North American Populus spp. so that the bronze poplar borer has not been associated with forest devas- tation. THE ASPEN ROOT GIRDLER. Harrison (1959) discovered what he thought to be a new species of Agrilus feeding on the roots of aspen suckers (Populus tremu- Carlson and Knight: Agrilus beetles 3 loides and P. grandidentata). The establishment of a chromosome number of 22 for this species seemed to indicate that it was more closely allied to anxius than to g. liragus (Nord, 1962). Later, Nord, Knight and Vogt (1965) discovered that the proper name for the root boring form is A. horni, a name which Fisher (1928) and Barter and Brown (1949) thought to be a synonym of anxius. Because of their uncertainty, however, they did not synonymize horni with anxius. A. horni has attained a modest reputation as a pest. Nord (1962) says that it caused severe damage to hybrid aspen out-plantings which were part of gen- etics experiments conducted by the Institute of Paper Chemistry in Wisconsin and Michigan. In 1964, the trees in some of the affected plots were destroyed and the tests were reinitiated. However, in 1964 they were still having trouble controlling horni! This, no doubt, was due to the inappropriateness of the con- trol techniques used, rather than to any factor in the beetle’s ecology which makes it difficult to control. Moreover, because of its ecological preferences, it is not likely that this insect will ever cause great injury to well-stocked nat- ural stands of aspen suckers that are growing on areas of moderate or good site quality. THE ALDER-BIRCH BORER. Fisher (1928) was uncertain of the status of A. pensus. He says ‘‘this species may only represent a color variety of anxius, but since nothing is known of its habits, it is best to retain it for the present, at least, as a valid species.’’ Even though he was unsure of the distinction be- tween pensus and anxius, he took the liberty of describing betulae, a synonym of pensus, from 3 specimens. Although there is really little question that pen- sus is actually a distinct species, its biology remained almost unknown prior to the studies here described. The ecology of pensus makes it rather innocuous, and it has attracted little attention. However, it appears to be an important fac- tor in the ecology of alders (Alnus spp.), which are pecan ee as wildlife cover in their lakeside and streamside habitat. TAXONOMY The Genus Agrilus SIZE AND DISTRIBUTION . Agrilus beetles are thought to feed only on angio- sperms (Fisher, 1928). Confinement to angiosperms could indicate that the genus arose in the Cretaceous or later, which seems to allow ample time for it to have attained the vast size and worldwide distribution it presently has. Schaefer (1949) said the genus included 2500 described species, and Obenberger (1957) claimed that in his collection alone, there were over 3000 described and undescribed forms. Obenberger (1959) believed that there are over 4000 species of Agrilus. The largest number of species occur in tropical or subtropical areas, but Agrilus is well-represented in temperate regions too. Fisher (1928) recog- nized 125 described forms from the North American fauna (North of Mexico), and a considerable number have been described since, mostly from the south- | Personal communication, 1964; D. Einspahr, Institute of Paper Chemistry, Rheinlander, Wisconsin. 4 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 western United States. SUBDIVISIONS. Obenberger (1957, 1959), with some rather specious rea- soning, tried to show that there is no way to divide Agrilus into acceptable sub- genera. He was, however, willing to admit that there are some recognizable Species groups. The economically important Eurasian ‘‘species’’! Agrilus viridis L., its many destructive ‘‘host races, ’’ and the lesser known North American A. po- litus Say together represent such a species group which is extremely difficult taxonomically. Both A. politus and A. arcuatus Say are classified by Fisher (1928) as being composed of several sympatric ‘“subspecies, ’’ but each actually represents a multi-species ‘‘complex. ”’ Frost (1912) recognized some of these ‘‘sibling species’? problems, and presented isa aes for their solution. He concluded by saying ‘‘the studies in the otiosus’? and anxius groups have been so far, rather dissappointing, due to the difficulty of getting series of both sexes. The only species that is at all abundant in this locality is otiosus, taken on oak leaves. The olivaceous vari- ety of acutipennis has been encountered quite often on oak, and bilineatus occurs in favorable places on oak sprouts, but in general the species turn up singly or in pairs, with aggravating slowness. Several very interesting problems are suggested by the material in hand, and more specimens from widely separated areas may present a solution.’’ In many groups of Agrilus, a similar situation still exists. The Agrilus anxius Group DEFINITION. Presumably broadening the limits of the Agrilus anxius group sensu Frost (1912), it is here defined as those species in which the adults have the carina of the pygidium projecting as a spine, (figs. 2, 3), and (2) have a sexual dimorphism in the structure of the last abdominal segment. The mar- ginal sclerite? of the hind segment is smooth and in males lies in a plane near- ly confluent with that formed by the sternite (fig. 33). In females the marginal sclerite is reflexed or beveled with respect to the sternite (fig. 32). In most of the species the first and/or second abdominal sternites are also sexually di- morphic. In males these sternites are often grooved medially (fig. 1), while fe- males never have such a groove. NOMENCLATURE Nord et al. (1965) have referred to the ‘‘Agrilus anxius complex.’’ The term complex carries a connotation of propinquity that is in- appropriate for a group that includes anxius, g. liragus, and horni, and we, anally includes a number of sympatric species (see, for example, Basurmanova, 1998). 2 eae Agrilus otiosus Say. “Those parts of the tergites termed the ‘‘vertical portions of the segments” by Fisher (1928), but they are not vertical on the hind segments and the term marginal sclerite seems to be a preferable alternative. Carlson and Knight: Agrilus beetles D therefore, will not apply it here. The Agrilus anxius group is comparable to what in many other families of insects would be recognized as a genus. Were the anxius group to be delineated from other Agrilus as either a subgenus or genus, either of the Agrilus synonyms Teres Harris (1829) or Uragrilus Seme- nov (1935) (=Epinagrilus Stepanov, 1954) could be applied. Because the limits of the anxius group are not ascertainable from the literature, it would be un- wise to use a formal name for it at this time. MORPHOLOGIC ANALYSIS . The taxonomic analysis of anxius group species is based primarily on adult morphology, but important information is also fur- nished by other developmental stages. The interpretation of morphologic char- acters is strongly influenced by studies of adult behavior which are discussed later, and some findings from studies of adult morphology are deferred to the Evolution section. Adults. The analysis of adult morphology was based primarily on collec- tions made in Michigan (Iron, Gogebic, Benzie, and Manistee Counties) and Wisconsin (Vilas Co.). Material in the collections of the University of Michigan Museum of Zoology (Ann Arbor, Michigan) the Entomology Research Institute of Canada (Ottawa, Ontario), Harvard University, Museum of Comparative Zoology (Cambridge, Massachusetts), the Philadelphia Academy of Sciences (Philadelphia, Pennsylvania) and the U. S. National Museum (Washington, D.C.) was also examined. The following types were studied: Agrilus betulae Fisher, Agrilus fulminans Fisher, Agrilus populi Fisher, Agrilus is horni Kerremans (=blanchardi Horn, name preoccupied), A Agrilus nevadensis Horn, Agrilus niveiventris Horn, Agrilus pensus Horn, and Agrilus gravis Leconte. On the basis of the study of the types and other material the following no- menclatural changes (including those made by Carlson [1969]) were called for and will be followed here: AGRILUS BROWNI, new species Agrilus browni, new species Agrilus lateralis, sensu Fisher (1928), in part. Fig. 1. Ventral aspect of Agrilus horni “, show- ing the median groove on abdominal sternites 1 and 2. 6 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 AGRILUS GRANULATUS Say Agrilus granulatus granulatus Say Buprestis granulata Say lata Say (1823) Agrilus grant granulatus populi Fisher; status assigned by Ss (1969) - Agrilus populi Fisher (1928) Agrilus Erichoca tpae Chamberlin (1929) Agrilus granulatus liragus Barter and Brown; status assigned by Carlson (1969) Agrilus liragus Barter and Brown (1949) AGRILUS PENSUS Horn Agrilus pensus Horn (1891) Agrilus betulae Fisher (1928); synonymy designated by Carlson (1969) AGRILUS QUADRIGUTTATUS Gory Agrilus quadriguttatus quadriguttatus Gory Agrilus quadriguttatus Dejean (1833, 1836), nomen nudum Agrilus quadriguttatus Gory (1841) Agrilus quadriguttatus niveiventris Horn; status assigned by Carlson (1969) Agrilus niveiventris Horn (1891) Agrilus nevadensis Horn (1891); synonymy designated by Carlson (1969) Agrilus quadriguttatus fulminans Fisher; status assigned by Carlson (1969) Agrilus fulminans Fisher (1928) AGRILUS RUFICOLLIS Fabricius Agrilus ruficollis Fabricius Buprestis ruficollis Fabricius (1787) Agrilus lateralis Say (1823), new synonymy Agrilus sayi Saunders (1871), name unnecessary (lateralis not preoccupied) Fisher (1922) reared an Agrilus sp. from bayberry (Myrica pensylvanica, = carolinensis) at Lyme, COBER CEI for which he used the name A. lateralis Say. The original description’ of lateralis (Say, 1823), however, fits A. rufi- collis Fabricius more closely than the species that was reared from Myrica. Forthat reason and because of the range limits apparent for it, the species which Fisher reared from Myrica is here given the name Agrilus brownin. sp. Say (1823) says the head of lateralis has a profoundly impressed line, which certainly is characteristic of ruficollis. A. browni has the front only weakly impressed. Say also notes that the length of lateralis is1/5 inch (about 5.1 mm), which is not exceptionally short for ruficollis, a smaller and more variable species than browni. On the other hand, specimens of browni as short as 5.1 mm must be extremely rare, as we have seen none approaching that size. I enordine to Fisher (1928) the lateralis type was lost, but, presumably, it was destroyed, as is the case for the majority of Say’s specimens. Carlson and Knight: Agrilus beetles 7 The only bonafide records for browni are from places very remote from the lateralis type locality. Say (1823) described lateralis from a specimen col- lected in Missouri, which then included the vast region west of the Mississippi known as the Louisiana Purchase, excluding what is now the state of Louisiana. Knull (1922) found browni in abundance on sweet ferns (Myrica asplenifolia, = Comptonia peregrina) in Pennsylvania, but no reliable records are known to us for any area west of that state. In spite of the examination of large amounts of sweet fern by P. C. Kennedy! and a modest amount of searching on that host plant by us, browni has not been encountered in Michigan. The strength of all the evidence leads us to suppress lateralis as a synonym of ruficollis. Fisher (1928) provides a good description of browni (under the name later- alis), and his ‘‘lateralis’’ neotype? is here designated as the type of browni. The species is named for Mr. William J. Brown?, who suggested that Fisher (1922, 1928)had misapplied the name lateralis. Fisher (1928) knew A. nevadensis Horn only from the type, and that speci- men appears to have been deformed during pupal development. Fisher (1928) says its scutellum is not carinate, but the carina appears to be present, al- though it is positioned abnormally on the deformed scutellum. The somewhat prolonged elytra have more sharply pointed apices and more coarsely rugulose surfaces than typical q. niveiventris, but there are no other structural differ- ences. The pigmentation of Some specimens of q. niveiventris from southern California is as dark as that of the A. nevadensis type, which was collected in Nevada. ee The following sections will be devoted primarily to anxius, pensus, horni, and g. liragus. The discussion of other members of the anxius group is de- ferred to a later section on evolution. Since Gory (1841) described anxius, other forms have been named, two of which are synonyms, while another was mistaken for a synonym. Agrilus gravis Leconte and A. torpidus Leconte were described in 1859, but Fisher (1928) placed them as synonyms of anxius. Agrilus blanchardi Horn (not Saunders) and A. pensus Horn were described in 1891. Because the former was a primary homonym, it was renamed A. horni by Kerremans (1900). Fisher (1928) thought that horni was a synonym of anxius. Knowing of no material besides the type Which could be identified as horni, he believed that it was merely an abnormal specimen. Being somewhat uncertain, however, he retained horni as valid. When Barter and Brown (1949) established that a poplar-feeding form and a birch-feeding form had been confused under the name anxius, they decided a He communication, 1967; P. C. Kennedy, North Central Forest Experiment Sta- tion, U. S. Forest Service, East Lansing, Michigan. en to Fisher (1928), this specimen is a male, which he collected on bayberry at Lyme, Connecticut and deposited in the U. S. National Museum collection. Uehncapliony Research Institute, Canada Department of Agriculture, Ottawa, Ontario. 8 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 to retain’ that name for the birch-feeding form. They suppressed torpidus as a synonym of anxius, but made no definite decision regarding the name gra- vis. The gravis type is a female that Barter and Brown (1949) could not identi- fy reliably. The name liragus was, therefore, given to the poplar feeding form with the distinct possibility of it later proving to be a synonym of gravis. There have been doubts that this was the proper thing to do. For example, the appli- cation of gravis by Blanchard (1889) for the species collected on poplars (not ‘“sprouts’’2) is cited by Nash et al. (1951) as a possible indication that gravis — was the proper name for the poplar feeder. _ Although the gravis type lacks the coppery sheen often found on the head and pronotum of anxius specimens, its absence is not a good indication of the iden- tity of the type. Vogt? pointed out that specimens of anxius from some parts of the Great Lakes Region are often colored more like g. liragus than are spec- imens from more eastern parts of the United States. Indeed, the coppery re- flections are often very indistinct in specimens from the Upper Peninsula of Michigan, which lies within the type locality recorded by Leconte (1859) as the ‘*Lake Superior Region. ’’ By relying on characteristics other than coloration and working with spec- imens collected in the Upper Peninsula of Michigan, we found that Agrilus gra- vis Leconte is really a synonym of anxius Gory, and therefore g. liragus Bar- ter and Brown is to be retained as valid The name gravis, as used in a num- ber of papers in the late 1800’s, refers to either anxius or g. liragus; assign- ment to one or the other depends upon the host designated, if any. References using the other anxius synonym, torpidus Leconte, are to be treated similarly, except for those which refer to willow as a host. The latter (e.g. Cook, 1890; Davis, 1891, 1892) refer to A. criddlei Frost, which makes galls on willows. The host of criddlei was recently established through rearing by Wong and Mc- Leod (1965). Barter and Brown (1949) concluded that anxius and g. liragus are separable only by reliance on male genitalia, coloration of the head and pronotum, and host plant identity. We cannot concur with them fully. As previously indica- ted, anxius specimens from some areas do not have very distinct coppery re- | flections on the pronotum, and this iridescence also seems to fade quite rapid- ly in Some museum specimens. Wetting the specimens with various relaxing agents is particularly destructive to the iridescence of these beetles. On the other hand, there are structural differences, besides male genitalic ones which, in combination, can be used for separating anxius and g. liragus. One of these, a difference in the form of the tarsal claws, was not noted by Barter and Brown (1949). The membraneous attachment of the tarsal claws of g. liragus appears to be either more flexible or looser than that of anxius. Consequently, the g. liragus tarsal claws are often clearly separated from the unguifer, making the Aathey, indicated the location of the anxius type as being unknown. “Blanchard thought what he had found on poplar sprouts was anxius. *Dersonal commununication, 1966; G. B. Vogt, U. S. Dept. Agr., A.R.S., Entomology Res. Div., Washington, D. C. Carlson and Knight: Agrilus beetles | 9 latter clearly visible. The claws of anxius are attached so that there is very little space between their bases and the unguifer. The unguifer, is therefore, more difficult to see. In addition, the claws of anxius are usually rather dis- tinctly swollen at their bases, and tend to obscure their attaching membrane which stretches across the opening in the last tarsomere, while in g. liragus the basal parts of the claws are usually not strongly swollen, and the membrane in the tarsomere opening is usually pti visible. The females of anxius and g. liragus differ in the shape of the projecting Spine on the pygidium, im, but the males do not. In g. liragus females the spine usually projects strongly (fig. 2) and is rather cylindrical, while in anxius fe- males it is usually shorter and more blunt (fig. 3), and its tip is often broad and elyptical in end view. A useful characteristic mentioned by Barter and Brown(1949), is the ten- dency for the elytral tips of g. liragus to be acute and strongly serrate (fig. Fig. 2. Dorsal surface of the pygidium of an Agrilus granulatus liragus 2 (X40). Fig. 3. Dorsal surface of the pygidium of an Agrilus anxius ? (X40). 10 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 A B C Fig. 4. Elytra of A. granulatus liragus o (A), A. anxius ¢ (B), and A. pensus ¢ (C). 4a), while those of anxius are more broadly rounded and not as strongly toothed (fig. 4b). By using both the coloration differences and the somewhat obscure and somewhat variable structural differences of the pygidial spine, tarsal claws, and elytra it is usually possible to distinguish the females of an- xius and g. liragus. iets The most reliable characters for separating males of anxius and g. lira- gus are those of the aedeagus. The parameres of anxius have broad ventral inner margins, and these margins together form a trough which is broad and shallow (fig. 5). Ventrally, the inner margins of the g. liragus parameres are usually narrow, together forming a trough which is narrow but deep in compar- ison with that of anxius (fig. 6). Nord et al. (1965) separate horni from anxius and g. liragus by utilizing characteristics of both the male and female genitalia. The ovipositor of horni (relative to elytron length) is significantly shorter than that of g. liragus or anxius. Nord et al. also pointed out that the average ovipositor length is less in g. liragus than in anxius, and in some cases the minimum length for g. lira- gus approaches the horni maximum. However, they found no overlap in ovi- positor length between g. liragus and horni. The male genitalia of these species correspond in length to those of the fe- males. In addition to being shorter, the aedeagus of horni differs from those of anxius and g. liragus in having parameres which are less dialated posterioral- ly, their margins being subparallel (fig. 7). Nord et al. (1965) say that ven- trally and subapically the inner margins of horni parameres are arcuate to subarcuate, while those of anxius and g. liragus are angulate to subangulate. Carlson and Knight: Agrilus beetles 11 Figs. 5-8. Ventral sides of the aedeagi of Agrilus anxius (5), A. granulatus liragus (6), A. horni (7), and A. pensus (8). The ventral trough formed by the parameres tends to be intermediate in form to the troughs of anxius and g. liragus, which are quite different from each other. Despite the confusion which has occurred regarding the identity of horni, it is really quite distinct from either anxius or g. liragus. Besides the geni- talic differences already noted, horni differs significantly from anxius and g. liragus in other ways. Most significant of all, perhaps, is the moderately dense pubescence on the marginal sclerites of the abdomen. As Horn (1891) points out, this pubescence appears all the more distinct because of the white pruinosity which is associated with it. Together, the pruninosity and white pubescence tend to form a white band which is very distinct on newly-collected horni specimens, but also quite noticeable on material that has been preserved for many years. The pubescence is usually dense enough to obscure the sculp- turing of all the marginal sclerites, except for the last one. These marginal sclerites are usually less pubescent in anxius and g. liragus, and the pubescence is usually not dense enough to obscure the sculpturing of the marginal sclerites. Moreover, the marginal sclerite of the second abdominal segment is either glabrous or noticeably less pubescent than those of the first and third segments in anxius and g. liragus. There are also distinct differences in the structure of the elytra. The tex- ture of horni elytra is very even, the disc being comparatively smooth, finely imbricate, and usually lacking a distinct longitudinal costa (fig. 9). The disc is usually more coarsely imbricate in anxius and g. liragus, and often has a 12 Contrib. Amer. Ent. Inst. vol. 4, no. 3, 1969 Pig... 9.) Agrilue horni ¢, longitudinal costa and a rather distinct depression just mesad of the costa and between the disc’s basal third and apical half. Frequently in g. liragus, and sometimes also in anxius (see o’, fig. 39), this depression is pruinose, the pruinosity forming a spot similar those on the elytra of g. granulatus, but less distinct. Upon close examination, the parts of the depression where the spot tends to form can usually be seen to have a small group of recumbent hairs which are alingned differently and sometimes longer than the surrounding ones. The pruinosity usually disappears in preserved specimens, but the faint spot formed by the unaligned hairs can often be detected. The recumbent pubescence on the elytra of horni is distributed rather e- venly over the surface and is also rather short and uniform in length. That of anxius and g. liragus is usually not uniform in distribution or length, and many of the hairs are longer than those of horni elytra. The structure of the head is also somewhat different in horni. The front is almost always rather flattened in profile. Sometimes in anxius and even less frequently in g. liragus, the front is somewhat flattened in din profile, but in these species it usually bulges, presenting a rounded profile. The form of Agrilus pensus is intermediate between that of A. anxius and A. quadriguttatus Gory, the latter being a willow feeding species. Barter and Brown (1949) say that the male genitalia of pensus are structured like those of g. liragus and g. populi, but this is not true. The pensus aedeagus (fig. 8) is somewhat variable, but more like that of anxius than that of g. liragus or g. populi. Barter and Brown also say that pensus lacks pubescent and prui- nose spots on the elytra, but while they are not found on specimens from Nova Scotia and New Brunswick they are quite apparent on some specimens from Michigan and are also found on the type of the pensus synonym betulae Fisher. As in A. quadriguttatus and subspecies the subapical maculation of n of the elytra is likely to be spread out along the elytral suture instead of concentrated in a small spot. Also as in A. quadriguttatus the elytra are likely to be angulate at the apex (fig. 4c), forming a broad notch, instead of acuminate as in A. eee s and subspecies, which have a narrower notch between the apices. Eggs. The eggs of pensus (Engel, 1968), anxius, g. liragus, and horni are creamy white immediately after they are deposited. They are flattened and Carlson and Knight: Agrilus beetles 13 oval in outline. There seem to be no interspecific differences in egg size, those of horni being about 1.2 mm long by 0.8 mm wide, according to Nord (1962). The eggs of anxius (and presumably g. liragus) are coated with a semi- transparent whitish substance immediately following deposition (Barter, 1957). The substance apparently acts as a cement and as a protectant for the soft egg shell. There is no evidence that horni applies such a substance to its eggs. Instead, the chorion of horni eggs appears to toughen as the larvae develop, and as a result they turn dark brown to black. The eggs of anxius and g. lira- gus darken like this only when parasitized. They develop grayish- -black streaks and then become completely black prior to emergence of the parasite adult (cf Barter, 1957, 1965). This led Nord et al. (1965) to believe that the darkening of horni eggs may have been associated with parasitism, but this is not the case. | Larvae. Benoit (1965) presents a complete description of g. liragus larvae, and gives the differences between these and the larvae of anxius. He says the larvae of these species are about 30 to 40 mm long and 2 to 3.5 mm wide . The body is white, except for the anal forceps and other minor sclerotized por- tions, which are ferruginous. The prothorax, which is wider than the rest of the body, has a median I-shaped sclerotized line on the tergum. A similar structure, which is only vaguely sclerotized, is located on the prosternum (Barter, 1957). The segments posterior to the prothorax are all about equal in width. Benoit (1965) says C-shaped spiracles are located on abdominal segments one through eight, and similar, but somewhat larger ones are found dorsolat- erally on the mesothorax. Actually, the sclerotized C-shaped structure is part of the spiracular valve, and the spiracular opening itself is located inter- nal to the transparent valve. Benoit (1965) states that the labial palpi are absent, but this is not true. Apparently he missed them because they were out of focus when his microscope was focused on the rest of labium (cf fig. 10 and his fig. 1B). Because of the nature of his samples, Benoit’s morphologic comparison of the larvae of anxius and g. liragus is of questionable value. He obtained his material from two localities, one for each species. This could hardly give one an adequate sample of intraspecific variation. Nevertheless, the species differences in the coloration of the anal forceps which Benoit has listed do appear to hold for larvae collected in Michigan as well as for those from Quebec. The anal forceps of anxius are described by Benoit (1965) as being reddish, with the line of sclerotization extending anteri- orally beyond the anal opening. He says the forcepsof g. liragus are more brownish, with the sclerotization ending abruptly just posterior to the anal o- pening. The larvae of pensus and horni tend to be intermediate to those of anxius and g. liragus in both the structure and coloration of the anal forceps. ~The characters Benoit (1965) used were worked out by Alexseev (1960). Among them was the location of a short macro-seta (‘‘un pied petite’’) located in relation to a small circular depression or dimple in the cuticle. The dim- 1 : Benoit (1965) does not specify the developmental stage of the larvae from which his mea- surements were taken. , 14 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 Fig. 10. Labium of Agrilus granulatus liragus lar- va (X234). Note that the right palp is completely out of focus as with both in fig. 1B of Benoit (1965). ples which are supposed to be associated with the pedalis structuralis (thoracic leg rudiments) are not evident on living larvae, and do not seem to be present with any regularity on specimens fixed in ‘‘XAAD’’ or by boiling. Consequent- ly, the dimple may be the result of shrinking or swelling of the cuticle or as- sociated tissues which may be related to the method of fixation and preserva- tion. Benoit (1965) killed and preserved his larvae in 70 percent alcohol. There also seems to be the possibility that the absence of the depressions may have been related to the stage of larval development. Most of the larvae we examined had assumed the bent-over position in the pupal chamber. Benoit failed to mention the developmental state of the larvae he examined. Benoit (1965) says there are also other marked differences between the larvae of anxius and g. liragus. As these require higher magnifications or more preparation, we did not examine Michigan material for these characters. Pupae. There is nothing particularly unusual about the pupae of these spec- ies. All are creamy-white at first, and gradually assume the adult coloration. The eyes are the first parts to darken, and the elytra darken last. It appears to be during the period of pupal development that much of the adult structural variation arises, and it may very well be as much due to variation in the con- ditions under which the pupae develop as it is to genotypic variation. GENERAL BIOLOGIES OF AGRILUS ANXIUS, A. PENSUS, A. HORNI, AND A. GRANULATUS LIRAGUS Hosts and Ranges THE BRONZE BIRCH BORER. A. anxius appears to be host specific to birches, Betula spp. Fisher (1928) lists the following birches as hosts for anxius: yellow birch, Betula lutea; paper birch, B. papyrifera; European white birch, Carlson and Knight: Agrilus beetles 15 B. alba (and varieties); gray birch, B. populifolia; and sweet birch, B. lenta. Barter and Brown (1949) also list B. fontinalis as a host for anxius. Hitchcock, et al. (1964) rank this birch as a variety of the western paper birch, B. oe dentalis, which is closely allied to B. papyrifera. Harlow and Harrar (1958), in fact, call the western paper birch B. papyrifera var. commutata. “Although it has been reared from numerous hosts, the principal hosts for anxius are the native paper birches, B. papyrifera and B. occidentalis. A. anxius is, then, primarily confined to northern forest types, but its range ex- tends somewhat south of the range of B. papyrifera in the eastern U. S., where it is found even at low altitudes as far south as Washington, D. C. It ranges west to Oregon and British Columbia. THE BRONZE POPLAR BORER. A. granulatus 1 iragus has been reared from several Populus species, although ‘its principal host is trembling aspen, Pop- ulus tremuloides. In addition, Barter (1965) lists the following as hosts for g. liragus: bigtooth aspen, P. grandidentata; eastern cottonwood, P. deltoides; balsam poplar, P. balsamifera ssp. balsamifera; and the black cottonwood, ¥% balsamifera ssp. p riehooarpe (Torr. & Gray) Brayshaw (1965). The known range of g. liragus is very similar to that of anxius, but g. lir- agus does not occur as far south as anxius does in the eastern U. S. The southern edge of the g. liragus range seems to comply more closely with the range of P. tremuloides than that of anxius does with the range of B. papyrifera. Just how far north either g. liragus or anxius occur is not known, ‘but both have been taken as far north as Edmonton, Alberta. The former, at least, should occur in interior Alaska. THE ASPEN ROOT GIRDLER. Trembling aspen is the principal horni host, but bigtooth aspen and balsam poplar are also infested. Infestation levels on balsam poplar are sometimes quite high. When seed stock of various species of poplars are transplanted in experi- mental orchard-like plots, horni can be particularly troublesome. Nord et al. (1965) found that under these conditions, horni readily infests the Eurasian species Populus alba and P. tremula. He found that various hybrid aspens were also infested. In nature, however, horni is probably confined mostly to species which reproduce by root suckering (i. e. aspens and balsam poplars). Although horni has not yet been found as far west as g. liragus, it is prob- ably not because the former has amore limited distribution, but is more likely a reflection of the fact that horni is not collected as readily as is g. liragus. There are specimens inthe U. S. National Museum from Massachusetts to Arizona, and as far north as Aweme, Manitoba. THE ALDER-BIRCH BORER. A. pensus has been reared from Alnus rugosa and collected on the foliage of this shrub. It has also been found in abundance on the foliage of green alder, Alnus crispa according to Barter and Brown, (1949). Fisher (1928) described betulae, a synonym of pensus, from speci- mens reared from river birch (Betula nigra) and Knull (1930) reared a large series from that host. The records of Blanchard (1889) for Agrilus granula- tus collected on alders refer to pensus | , not to quadriguttatus as Fisher (1928) suggests. Records for pensus on Ostrya virginiana are of questionable sig- nificance. One specimen collected by C. A. Frost (Monmouth, Maine; June 23, 1910) is labeled ‘‘ironwood’’ and three by Geo. M. Greene (Philadelphia, 1 Horn (1891) described pensus from the series taken by Blanchard (1889). 16 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 Pennsylvania; May 24, 1900) are labeled ‘‘hophornbean.’’! Not only is it rath- er doubtful that pensus infests Ostrya, but it is entirely possible that alders or young birches were mistaken for Ostrya. Ironwood, at least, is an unlike- ly host because its ecology is so very different from that of alders and river birch. A. pensus has been collected as far west as Aweme, in southern Man- itoba, and in the East occurs as far north as mid-New Brunswick (Barter and Brown, 1949). Manee (1913) recorded Agrilus granulatus on ‘‘black alder’’ at Southern Pines, North Carolina. As it was collected with Eupristocerus cogitans Weber (an agriline species that infests alders), the record should be considered to refer to Agrilus pensus. Life Cycles A. anxius and A. granulatus liragus have a two-year life cycle under some conditions, but a one-year life cycle when other conditions prevail. A. horni and A. pensus, however, seem to have obligatory two-year life cycles. ANXIUS AND G. LIRAGUS. When the eggs of anxius and g. liragus are de- posited on hosts that are dying, severely injured trees or blowdowns, for ex- ample, the larvae often construct the pupal chamber during the same season that the eggs were laid. On the other hand, when oviposition occurs on more vigorous hosts, a two year life cycle results (Balch and Prebble, 1940; Ander- son, 1942; Barter, 1957, 1965). Nonetheless, the larvae of these species cannot survive in healthy trees, and successful larval development is always dependent upon the host being in a weakened condition because of drought, re- peated unsuccessful borer attacks, or other injuries (Barter, 1957, 1965). PENSUS. The habits of A. pensus have not been studied sufficiently to es- tablish the usual condition of its host at the time of oviposition. We have never seen larval galleries that would indicate a life cycle of less than two years for pensus. Furthermore, the larvae appear to overwinter primarily in two stages. Mature larvae, of course, overwinter in the pupal chamber, and early instar larvae are found overwintering deep within the xylem. Blanchard (1889) says pensus ‘‘breeds in and frequents stems of partly dead alders, but does not appear on the foliage very much.’’ Knull (1950) says it breeds in ‘‘unhealthy’’ river birches, but mentions nothing about the size of the trees in- fested. Fisher (1928) says the pupal cells are common in the stems of young river birches that have died, and we have similarly found that the adult emer- gence holes are common on dead alders. However, numerous speckled alders were examined for the presence of pensus larvae, and those which harbor im- mature larvae are invariably alive, and many of those infested with mature larvae or later stages have not succumbed completely. Because the larvae al- ways seem to require more than one year to complete development, the adults probably never oviposit on hosts imminently near death. Only one attempt was made to attract pensus with cut stems of alders. Although there was a pensus infestation in the area, none were attracted. Ovi- position on such material would be detrimental for this species, because early death would be inevitable for the larvae. The bark of Alnus rugosa is so thin that the phloem would become moribund and dessicated much too quickly to Noseinens in U. S. National Museum, Washington, D. C. Carlson and Knight: Agrilus beetles 17 permit larval development. However, it seems probable that even large, de- cisively injured or felled river birches are not attractive to pensus. HORNI. Nord (1962) found no evidence of anything less than a two-year life cycle for horni, and those findings are supported by our observations. By examining suckers containing mature larvae it is often possible to establish when the first instar gallery was formed. First instar larvae feed in the phloem! while moving down the stem toward the root, but in no instance was a first in- star gallery located in the phloem of suckers infested with mature larvae. When it was not obliterated by the larva when it re-entered the stem, the dark spot indicating the position of the first instar gallery was invariably found between the outer two rings of xylem. Moreover, out of 60 larvae introduced into suckers of various sizes, 5 lar- vae completed their development during the second summer. One larva ma- tured in the laboratory during the third summer, but no larvae matured during the first season. Nord (1962) found that the total length of horni galleries varied between 40 and 90 inches. The maximum length of the galleries of anxius and g. liragus which mature in two years is about 52 inches. The galleries of anxius or g. liragus larvae completing their development in one year are usually less than half that long (cf Barter, 1957; Anderson, 1944). Thus the evidence is over- whelmingly against the occurrence of a one-year life cycle for horni. Two years are usually required for horni development to be completed, but occa- Sionally it may take three years. Developmental Patterns HATCHING OF EGGS. Barter (1957, 1965) found that incubation of anxius and g. liragus eggs requires about two weeks. The same is true for horni, and is probably generally true of many Agrilus. When embryonic development is complete, the first larval instar bores out of the egg and into the bark of the host, leaving the egg shell packed with frass. LARVAL FEEDING PERIOD. Although larval behavior differs among Agrilus species, certain generalities can be made. As with most Buprestidae, the lar- vae never bore exit holes for their frass, but instead pack it tightly into the gallery behind them. Heering (1956) gives a very detailed account of the larval behavior of Agri- lus viridis. Some aspects of its behavior seem very similar to those of anxius or g. liragus, and other aspects seem typical for Agrilus in general. The viri- dis larvae are pressed tightly into the gallery. According to Heering this allows the microscopic spinules on the lateral muscular ridges to grip the sides of the gallery. The larvae move forward by constricting and stretching the segments telescopically. Movement of the parts of the body besides the head and anal for- ceps is entirely restricted by the sides of the gallery. All the tissues macerated by the larvae pass through the digestive tract. As the frass passes out the anus, it is clasped between the anal forceps. With these the larvae pack the frass tightly into the gallery behind them (Heering, lord et al. (1965) said that during the first stadium the larvae feed in the cortex, but this is the position of the gallery one year after it is formed. 18 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 1956). It would seem necessary that the frass be packed at least enough to make its density as great as that of the tissues on which the larvae feed. The Bronze Birch and Bronze Poplar Borers. Barter (1957, 1965) says that after they. bore out of the egg, the larvae move directly through the bark to the cambial'’ region, but Anderson (1944) found that some larvae bore as much as 10 inches in the bark while passing obliquely to the cambial region. These species mine chiefly in the phloem, but the xylem is also scored. An- derson (1944) found that after reaching the cambial region, the larvae feed for an average of 1.3, 3.9, 7.1, and 13.0 inches in each of the four larval stadia preceding the molt to the fifth instar, after which the pupal chamber is formed. This indicates that there is a correlation between larval size and the amount of feeding done in any stadium. In the trees from which Anderson accumulated these data the points at which molting occurred were marked by departures of the gallery into the xylem. In hosts of a very decadent nature the larvae usu- ally do not bore into the xylem before molting. On the other hand, in the more vigorous hosts which can be successfully infested, there are likely to be more xylem departures than molts. Apparently, these xylem departures are elic- ited by the unfavorable conditions occurring when host vigor is comparatively high. Host condition also has a bearing on other aspects of gallery formation. In severely weakened or very decadent hosts larval galleries usually show no distinct pattern. Under these conditions, the larvae follow a course suggesting that they feed on the freshest phloem they encounter (Barter, 1957). These galleries can be fairly straight, but they usually meander without any pattern in these kinds of hosts. In more vigorous hosts a zig-zag or sinuate gallery pattern is the rule. Under these conditions anxius makes a gallery which progresses in the direc- tion tended by the phloem sieve elements or the xylem vessels, but weaves back and forth across their grain forming a gallery with continually expanding hyberbolic links (figs. 11, 16). Under similar conditions the successive links Fig. 11. Larval ely of Agrilus anxius (in Betula papyrifera) that has healed over, indicating that it was made under conditions of high host resistance (from Anderson, 1944). eine authors suggest that these insects bore in the cambium. The cambium is extreme- ly thin (strictly defined, a single cell thick; cf Esau, 1953) and, therefore, it is inaccur- ate to say that these insects bore in the cambium. Although the cambium may be con- sumed as an insignificant part of the nutriment, food value is more likely to be derived from its derivatives, particularly the phloem and perhaps the newly-formed xylem. Carlson and Knight: Agrilus beetles 19 of the g. liragus gallery are much closer together, often lying adjacent to each other (fig. 12). The attempt to explain these gallery patterns will be deferred to the part of this section which deals with host resistance. The Alder-Birch Borer. The larvae of A. pensus boring in the stems of Alnus r rugosa iosa often form galleries without any regular pattern. Sometimes, however, the gallery is sinuate like those of anxius, particularly those parts of the pensus gallery which do not lie deeply in the wood. Xylem departures are more frequent with pensus larvae, and sometimes they bore through the wood from one Side of the stem to the other, a behavioral trait noted by Bar- ter (1957) for anxius when it bores in branches. According to Larsen (1901) anxius larvae make spiral galleries in branch- es. This has never been reported for g. liragus, and pensus does not form Spiral galleries, even when it bores in alder branches only about an inch in diameter. Only one case was encountered in which several pensus larvae had com- pletely girdled the phloem! , and this was only during the latter stages of lar- val development. Very little is known about the larval behavior of this spec- ies, but it seems clear that if the host dies during the earlier stages of lar- val development, they would be unable to complete their development (cf page 16). Perhaps for this reason, mass attack is not the habit of this insect. The Aspen Root Girdler. Some aspects of A. horni development seem to differ radically from what is characteristic of anxius, pensus, and g. liragus. In horni and often in the others, eclosion from ‘the eg egg is followed by direct entry into the phloem, but the first instar gallery is otherwise quite different in horni. The mechanism triggering the first molt of horni larvae appears to Fig. 12. Larval galleries of Agrilus granulatus (from area of in- tergradation ofthe nominate subspecies with g. liragus [see p. 85], but typical for the latter, at least). Some had healed over, indicat- ing high host resistance (courtesy of R. E. Stevenson, Forest Biol- ogist, Canada Dept. For. & Rural Dev. , Calgary, Alberta). OP on ieke girdling may be more prevalent under some conditions than others. At Mall- ard Creek in Iron Co., Michigan, a moderately large pensus population build-up occurred one or two years after a stand of alders had been flooded by a beaver dam. In this area, alders that had been completely girdled by pensus were not rare (personal communication, 1968; R. P. Engel, University of Michigan, Ann Arbor). 20) Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 be different, because the first instar galleries of this species are comparative- ly long. Prior to molting the first instar larvae of anxius and g. liragus bore an average of 1.3 inches after entering the phloem, although the gallery some- times extends as much as 10 inches in the bark before it reaches the phloem (Anderson, 1944). In suckers recently killed by horni what appears to be the first instar gallery may extend out into a root for as much as 42 inches, and the total length of the first instar gallery usually is greater than 12 inches. Whether or not the first molt of the horni larvae occurs during the first sea- son of development is unknown. No larvae have been found at this stage during the winter or spring. Presumably, they have been overlooked because of their small size (about 2 mm long in the first stadium). Brooks (1914), presents an excellent discussion of the larval behavior of Agrilus vittaticollis Randall, which probably is a distant relative of horni (see fig. 44). He says that upon leaving the egg, the vittaticollis larva bores in the ‘‘cambium’’ (meaning cambial region) progressing down the trunk and into the root for a distance of 6 to 12 inches before boring into solid wood. For most of its length the gallery in the phloem is straight, generally following the grain of the wood. However, it sometimes zig-zags across the grain af- ter leaving the egg, and invariably spirals once or twice in the phloem of the root before entering the xylem. The larvae of horni behave similarly, but proceed with the grain until they enter the root. Just after reaching the root they are apt to make one or two Spirals in the phloem, and often make others before they begin boring princi- pally in the xylem. This behavior varies and may be dependent upon the size of the root and the condition of the host. Sometimes the horni larvae do not bore in the xylem until they turn back toward the stem. In other instances they spiral in the xylem while still pro- gressing away from the stem. Root diameter seems to be the factor deter- mining where the larvae change the feeding direction. The turn is apparently elicited when the larvae reach a point where the diameter of the root they oc- cupy is so small that they completely destroy the xylem. In aspens this is apt to occur where the root diameter is about 1/8 to 3/16 of an inch in diameter. The larvae may derive the stimulus eliciting the turn toward the stem when severing of the xylem results in different pressures on each side of the breach. While transpiration should make the pressure on the proximal side negative, any pressure in the distal part should be positive. The turning point may be as much as 42 inches or more from the stem, but if the roots are short the larva may reverse its feeding direction only 6 inches from the stem. Instances in which the turning point is close to the stem are rare because the roots of aspen suckers are usually rather long. The phloem in the roots of young aspen suckers is much thinner than that in aspen or birch stems suitable for infestation by g. liragus and anxius, re- spectively. The difference relates to the age-dependent accruement of inac- tive phloem. Although horni galleries lie principally within the xylem, it could hardly be otherwise. It appears that they consume the phloem too, and of all they ingest, the phloem probably provides most of what is assimilated. After horni larvae re-enter the stem they sometimes bore a rather loose- ly spiralled or almost straight gallery which departs from the cambial region. Nord (1962), however, notes that the larvae always tunnel in the cambial re- gion long enough to girdle the phloem. The root damage inflicted by horni larvae rarely seems to cause the death of suckers, because foliar discolora- tion occurs after the larvae return to the stem. Zahner and Debyle (1962) Carlson and Knight: Agrilus beetles 21 found that pruning only one half of the parent root of aspen suckers causes lit- tle mortality, even after two years. Consequently, horni-induced sucker mor- tality must be considered to result mostly from girdling of the stem. The extent to which horni larvae feed in the cambial region after returning to the stem may be related to sucker vigor (Nord, 1962). Stager (1923) found that Agrilus aurichalceus Redtenbacher (= rubicola communis Obenberger) boring in bramble (Rubus spp.) stems increases or decreases the distance be- tween spirals such that the stem does not die until the period of pupal maturity. Apparently, the death of the stem is a crucial factor, because Stager says the pupae require a dry chamber. This seems to be true for species in the anxius group, as well. One might, therefore, expect variations in the position of horni pupal chambers to be at least partially related to moisture conditions in the stem at the time of its construction. Their pupal chambers are located an average of 4.8 inches above ground, but the position varies from a height of 2 inches to 14 inches. (Nord et al., 1965). POST-FEEDING LARVAL PERIOD Formation of the Pupal Chamber. Brooks (1914) made a detailed analysis of pupal chamber formation by A. vittaticollis. The vittaticollis larvae ex- tend a burrow of the usual dimensions through to the bark at the upper end of the gallery. They then recede about 1/4 inch and begin eating wood from the sides of the gallery. As this proceeds, their feeding is directed backward along the ventral side of the body, thereby doubling the gallery diameter. The two ends of the body are, as a consequence, appressed closely together, and the head and anus soon lie side by side at the bottom of the pupal chamber. The chamber is then complete, but the larvae continue to move forward until the head and anus are at its upper end. The larvae of all Agrilus spp. probably construct the pupal chamber sim- ilarly. However, when the layers of birch bark are gradually peeled off over anxius pupal chambers, a small loop in the gallery is found at the upper end of the chamber (exit point for imago). The loop apparently marks the point at which the larva turned in the bark and re-entered the xylem. It appears to be peculiar to anxius, and may somehow relate to the peculiarities of birch bark. The loop is not evident at the exit points constructed by g. liragus larvae, and horni and pensus, at least, must double back in the manner Brooks (1914) has observed for vittaticollis. Shortening of the larva may be initiated earlier in most of the other spec- ies of the anxius group than it is in vittaticollis. Whereas vittaticollis initially settles in the pupal chamber with the body folded in half, the larvae of other species assume a position in which the body is folded between the second and third abdominal segments. While vittaticollis has both head and anus at the upper end of the chamber, the other species have the folded portion of the body at the upper end, the anus at the lower end, and the head pointed downward ad- jacent to the abdomen (in standing trees). This would indicate that instead of stopping where vittaticollis does, with head and anus at the upper end of the gallery, they move around farther. An alternative possibility is that after boring to the outer layers of bark, they merely turn back and feed along the ventral side of the body until the gallery width is doubled, then gradually re- cede or shrink to the overwintering position. This seems unlikely, however, as the exit point is plugged with frass. Unless these Agrilus have acquired the ability to pack frass with the mouth parts and head, it must be assumed that the anus, at some time, is at the upper end of the pupal chamber. Brooks 22 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 (1914) implies that it is the fourth instar vittaticollis larva which constructs the pupal chamber, which may explain the difference in the folding position for the abdomen in this species. The molt to the fifth instar occurs before forma- tion of the pupal chamber in pensus, anxius, horni, and g. liragus, and the fifth instar larvae are noticeably shorter and more stout than fourth instar lar- vae. Diapause. Barter (1947) has shown that it is necessary for anxius larvae to reach the fifth instar before overwintering in order for pupation to occur the following spring. He says that subjection of fifth instar larva in the pupal cham- ber to subfreezing temperatures is necessary for the continuation of develop- ment. In early October 1966, some sections of a small birch tree infested with anxius were collected near Manistee, Michigan, and stored indoors with the intention of removing the larvae from them at leisure. In November part of them were debarked, and none of the larvae had shrunk to the prepupal form. In early December, however, adults emerged from these logs, and a few more issued in late December. Some larvae, on the other hand, did not enter the prepupal stage, suggesting that only some of them had been exposed to low temperatures in the proper stage and for a sufficient duration before they were brought indoors. This is in agreement with recent theories which, according to Wigglesworth (1964), suggest that diapause involves some phase of develop- ment that can proceed only at low temperatures. While the transition from the overwintering stage to the prepupal stage may be possible only after some exposure to low temperatures, the transition it- self appears to occur only when temperatures are above Rie and probably considerably higher. In Michigan the transition of anxius, g. liragus, and hor- ni larvae to the prepupal stage occurs during the spring. In Benzie Co. the” transition of liragus larvae, in some cases, occurs before late April, but usu- ally is later. In Iron Co. some horni larvae do not go into the prepupal stage until early June. PUPAL TRANSITION. In Iron Co. the transition from the prepupa to the pupa usually occurs in May and early June for anxius, g. liragus, and horni. There is no delay in development when the pupal stage is reached, and soon after the all-white pupa forms it begins to take on some of the adult coloration. The eyes are the first parts to darken, and the mandibles darken next (Barter, 1957). The elytra remain white until most of the other parts of the body have darkened, and do not come to rest in the adult position until rather late in pu- pal development. Very likely, this is the reason that the wings are deformed more frequently than other parts in rearing these beetles. When drying of the wood around the pupal chamber is too rapid, the elytra may not take shape properly, after uncurling from the sides of the abdomen. This usually results in a poor fit of the elytra along the suture, causing a gap which exposes the membraneous wings. Sometimes the deformities are more extreme. ADULT EMERGENCE Study Procedure. Eclosion of the adults from the pupal chamber was fol- lowed in several ways. Unfortunately, no good field emergence data were ob- tained for g. liragus, and no attempts were made to get records of field emer- gence for pens In 1964 g. liragus emergence began about May 30, which was considerably earlier than was expected. Consequently, there was not sufficient time to lo- cate enough pupal chambers for checking field emergence adequately. Some data were obtained by placing small triangular screen cages (fig. 13) over the Carlson and Knight: Agrilus beetles 23 exit points that were exposed by cutting away the outer bark layers. The same method was employed in 1965, but because emergence was considerably delayed the results were extremely poor. Dessication of the exposed bark layers and wood prior to the beginning of the emergence period caused high mortality of the pupae and young adults. The same type of cage worked well for analyzing field emergence of anxius. To expose the exit points for anxius adults, only a few outer layers of birch bark had to be removed. Consequently, the rate of dessication of inner bark and wood probably differed insignificantly from that of infested trees not trea- ted in this way. A. anxius emerged normally from caged trees in 1964 and 1965. In 1964 the cages were checked intermittently, only often enough to roughly establish the beginning, end and peak period of emergence. In 1965 they were checked daily, except on days when emergence was unlikely. In 1964 the emergence of horni adults was followed in the field by examin- ing some infested suckers marked with flagging. Although the plot was checked only every other day, this method proved to be rather inefficient. Hence, dif- ferent methods were used in 1965. The basal portions (sticks) of 123 infested P. tremuloides stems were placed in a clearing at the University of Michigan Forestry Camp near Beechwood, where they received full sunlight during the middle of the day. The ends of the sticks were waxed, and the pupal chambers were then located by comparing the sticks with their X-ray! negatives. Tubes of plastic screening were positioned over the segments of the sticks harboring the pupal chambers, and the ends of the tubes were constricted by stapling or taping. The sticks were held in an upright position near the ground by cords strung between stakes (fig. 14). They were checked daily. It is probably safe to say that in most parts of their ranges, peak emer- gence for anxius, g. liragus, and horni occurs during June, but even within a given region there can be significant year-to-year variations in the emer- gence pattern. The variations in emergence periods relate to weather varia- tions. Davis and Raghuvir (1964) compared deviations from long term mean air temperatures with emergence data for A. aurichalceus. The temperature data used were those for the month of March, April, and May. Walton (1951) Fig. 13. Small emergence cages stapled over exit points for Agrilus anxius adults. 1 The X-ray machine and techniques used are discussed by Knight and Albertin (1966). 24 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 Fig. 14. Plot at Camp Filibert Roth near Beechwood, Michigan, where the emergence of Agrilus horni was studied under semi-natural conditions by stringing the infested sticks between stakes to hold them vertically. employed the same technique for studying the emergence of A. ruficollis Fabri- cius for successive years. Emergence appears to be sufficiently well corre- lated with monthly temperature data to make the technique useful. Results and Discussion. Table 1 shows the deviations from long term monthly means for Stambaugh, Michigan for March, April and May in 1964 and 1965 (U. S. Weather Bureau, 1964).. The average deviation for 1964 was +0.97 degrees, while that for 1965 was -1.23 degrees. Emergence data for these two years reflect the differences. In 1964 anxius emergence began in the Stambaugh vicinity about June 4 but not until June 14 in 1965. The difference between peak emergence times was even greater, being about June 7 or 9 in 1964, and between June 20 and 23 in 1965. In 1964 horni emergence began about June 7, but was delayed until a- bout June 20 in 1965. Peak emergence was between June 9 and June 15 in 1964, while it occurred between June 21 and June 28 in 1965 (fig. 15). The 1964 emergence periods of horni and anxius did not extend past the end of June, but in 1965 both species continued emerging into July. Surpris- ingly, one anxius adult was found alive in an emergence cage on August 4, but with this unusual exception, emergence of this species ended by July 7. A. horni emergence ended on July 15, making the duration of the 1965 emer- gence period about equal to that for 1964. Table 1. Average deviation from long-term mean air temperature (F) for Stambaugh, Michigan. eich 1964 co 1965 March “eae ~O. 6 April +13 “2. b May ab ih +4.0 Mean +0.97 -1.23 25 Carlson and Knight: Agrilus beetles ‘arlojoq skep Z IO [ uUeY} 91OW poSi1oWIN pey }I JI peep useq sAvY PTNOM pue F 4SsNsny UO oSvod sOUSS.IOWIS US UI BAITS PUNO} svmM }[Npe sNTxue ouGO (Zz) ‘UMOUYyUN ore soyep ooues -IOULS OTJIoeds puv peYxdeYO oq JOU P[NOd j4oTd 9} [BAIOJUL STU TOF ‘POST ‘ouNL JO YI 6Z 94} YSno1Y} YIOT OY} Wor periMmd90 SooUdSIOWO TULOY SATA (T) “SUIMOT[OF 94} OJON ‘“UBSTIYOT|! ‘*OD UoA ul ‘dds sn{lisV Jo soyep soueS10UIN YJNpY “CT ‘BIW i ec Pe ee 0 Gg6l ‘SnSeaT] SnyejnueIs SniIwy OI © re Aed ied Suisi9sw gy seljeog jo 19quINN G96 ‘SnIxue SnTIIsV 0 bo S Ui) & oD) POEL ‘TULOY SNTIIsy OL Pete wor tee Poe eS Pee ke 6 Ce eS ee ETS ORS PS eee Tee eh 9 eg ATOL ANAL 26 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 Barter (1949) found that the mean emergence dates for g. liragus preced- ed those for anxius by an average of 12.7 days. Although data obtained for g. liragus inour study were rather poor, it is evident that the differences in time of emergence are not as great in Michigan. A. g. liragus began emerg- ing about 7 or 8 days before anxius in 1964, but the peak emergence periods were not as far apart. Anderson (1944) reared both anxius and g. liragus from caged aspen and birch logs in Minnesota. He considered the possibility that he was working with two ‘‘host races, ’’ so one might assume that he would have noticed any significant difference in emergence times. In combining the emergence data for both anxius and g. liragus, Anderson found that emergence began on May 29 and ended on July 2, the peak occurring on June 10. A difference in peak emergence times for g. liragus and anxius would ap- pear to have no more real significance than any other differences between these species, and may be subject to the same kinds of geographic variation. These beetles are, in any case, actively mating and laying eggs during time spans which are nearly coextensive. Population Regulating Factors The factors which are usually considered important in the regulation of insect populations are parasites, predators, and weather conditions. There have been many semantic arguments devoted to the question of whether or not factors which cannot take an increasing toll on an organism as its density in- creases can be as important as factors that can increase their toll with density. Because factors like weather conditions have no way of responding to density, it is usually assumed that they fall in the former category. More often than not, though, weather conditions are clearly a part of the climatic scheme to which a species becomes adapted. There may be some variations of weather which occur so infrequently that they are not likely to bring about genetic adapt- ation in the species affected. Such catastrophic factors are here considered to be of secondary importance as population regulating factors; a species must, in the long run, be part of a density dependent controlling system. The factors likely to be of major importance in controlling the Agrilus being considered here, are parasites, predators, and host resistance. HOST CONDITION. Host resistance can be regarded as an indirect effect of weather conditions, but it is probably more valid to consider it to bea manifestation of the more or less balanced relationship between the host and | its phytophagous parasite, and a manifestation of the physiological condition of the host. The physiological condition of trees is most directly related to their age and site quality. By loosely applying these two variables one could explain most of the interactions which occur between Agrilus spp. and their hosts but the balance of the interrelationship is also influenced by parasites and predators, both phytophagous and entomophagous. The Bronze Birch and Bronze Poplar Borers. Anderson (1944) says the larvae of anxius or g. liragus require living ‘‘cambium’’ if they are to com- plete their development successfully. Inner bark tissues which have died and turned brown are unsuitable for larval development. Barter (1957) found that some first instar anxius larvae may bore 13 to 18 inches without molting in moribund phloem. Those that are able to molt bore about twice as far (6 inch- es) as larvae feeding in phloem which has not turned brown. The evidence Carlson and Knight: Agrilus beetles 27 seems to indicate that these larvae require nutrients associated with living phloem and that these essential nutrients decompose when the phloem cells die. | | Swaine (1918) and Balch and Prebble (1940) reported finding anxius attack- ing and killing many healthy, mature birches in Canadian forests. Nash et al. (1951) reported that during times of high bronze birch borer population levels, healthy trees are attacked and killed. Different opinions were held by Spauld- ing and MacAloney (1931) and Hall (1933), who believed that the borer attacks birches because they are ‘‘dying.’’Barter (1957) suggests that the borer may appear to be more aggressive under outbreak conditions, but that it otherwise plays only a secondary role. He and Anderson (1944) present evidence show- ing that the beetles are usually unable to successfully attack healthy trees un- less they are exposed to repeated unsuccessful borer attacks in successive years. Of course, after such repeated attacks, the trees could not be con- sidered healthy, and persistent attacks of this sort could only occur during extended outbreaks. Perhaps unsuccessful attacks indicate that the number of susceptible trees is limited, or that ‘‘spill-over’’ occurs from highly attractive trees onto less attractive ones adjacent. It is apparently only when many trees are simultaneously predisposed to infestation that the bronze birch borer becomes epidemic. A. granulatus liragus apparently does not often reach the outbreak state, but Barter (1965) and Barter and Cameron (1955) found that following repeated defoliation by the forest tent caterpillar, Malacosoma disstria Hber., aspens are predisposed to g. liragus infestation. Under such conditions g. liragus can cause deterior- ation and death of hosts it attacks either on branches or the main stem. Arru (1962) discusses the factors which predispose newly transplanted poplars in northern Italy to attacks of Agrilus suvorovi Obenberger (=viridis populnea Schaeffer). Problems with this insect arose only after poplars were planted on soils which were not as well suited to them as the deep friable soils to which poplar cultures were formerly confined. Arru believes that the factors which make the transplants susceptible to infestation by the beetle are those preventing resumption of growth following planting. The most important of them is moisture stress, which is likely to occur when poplars are transplanted on sandy or pebbly soils, on embankments, or in compact, poorly aerated soil. Anderson (1944) treated aspens and birches in a number of ways to deter- mine the severity of injury, if any, necessary to predispose them to the attacks of g. liragus and anxius'. He made some interesting observations on brood development in 6 aspens and 6 birches which were girdled in September, 1940. All of these trees, except one of the aspens, remained alive throughout the summer of 1941, when they became infested with g. liragus and anxius. Brood density below the girdles in the aspens averaged 1.2 per sq ft (52 sq ft exam- ined), while above the girdles there were only 0.1 larvae per sq ft (88 sq ft examined). Similar results were obtained for the birches. Below the girdles all the living borers were either pupae or adults, but only larvae were found above the girdles during the spring of 1942. All of the trees produced a small annual ring above the girdle during the summer that they were attacked by Agrilus. No growth rings were produced below the girdles in any of these trees, however. When Anderson (1944) debarked the girdled trees he found that the bark Both taxa were still lumped under the name anxius at that time. 28 Contrib. Amer. Ent. Inst., vol. 4, no.3, 1969 moisture content below the girdles was nearly 2 times that above. He assumed this to be a reflection of conditions that had existed during the previous summer. This assumption is incorrect because girdling trees by cutting through the phloem causes starvation and death of the roots. After the roots of a girdled tree die, water absorption ceases and the rest of the tree probably dies slow- ly from dessication (Kramer and Kozlowski, 1960). Because the phloem above the girdle continues to receive small amounts of carbohydrates, while that be- low gets none, the former should maintain a higher moisture content. It is the presence of solutes (sugars, etc.) in the phloem which allows its turgidity to be maintained. The diffusion pressure deficit of phloem sap offsets the effects of negative pressure or tension in the xylem. Therefore, the high bark moisture contents Anderson found below the girdles in the spring of 1942 were possible only because the bark had absorbed moisture during the winter. Hence, he has used a false premise in asserting that the larval mortality in trees with a proliferating cambium is not related to bark moisture content. Heering (1956) found a relationship between bark moisture content and the susceptibility of beeches (Fagus sylvatica) to attack by Agrilus viridis. However, he claims that in some cases callose formation becomes the impor- tant host resistance factor, especially in trees so extensively dessicated that defensive sap flow is impossible. Because callose is a polysaccharide con- sisting of linked glucose units (Kessler, 1958), its formation involves a proc- ess much like normal growth. The growth of trees is probably controlled more by water availability than by any other environmental factor (Kozlowski, 1964), and deficits in water availability can cause physiological conditions which override growth regulatory processes (Zahner and Donnelly, 1967). It is, therefore, difficult tosee how severely dessicated trees could form callose fast enough to pinch off continuously feeding larvae. The most logical explana- tion is that when Heering thought larval mortality was attributable to callose formation, the larvae had, infact, been overwhelmed by phloem exudates, and were afterward being overgrown by callose. Even his own observations sup- port this contention. He found that in very moist, vigorous bark, the virdis larvae construct vent holes (‘‘Rindensaft-Abflusskanalen’’) which prevent sap from accumulating in the gallery by allowing it to drain off to the outside. The very fact that the larvae have time enough to cease forward movement and push the anterior part of the body through the bark, suggests that wound cal- lose formation could not be rapid enough to kill them. Heering (1956) found that the moisture content of bark infested with first instar larvae is almost as high as that of sound bark. This could indicate that bark which appears to be sound may really have low vigor, or instead, that his technique is too crude for measuring the essential differences between susceptible bark and bark resistant to infestation. He removed his bark sam- ples from the trees, placed them in glasers’ putty, and analyzed their mois- ture content in the laboratory. His index of tree resistance was bark mois- ture content expressed as a percentage of its total dry weight. This measure of host resistance is not precise because the active phloem constitutes a very small fraction of the total bark mass. Critical differences in its turgidity would be masked by variations in other bark factors, such as the amount of accumulated cork or innactive phloem. Perhaps a refinement of the technique, such as that used by Bier (1964) would give better results. However, phloem pressure would probably be a better measure of active phloem borer resistance for the majority of woody angiosperms. Some of the angiosperms have gum ducts and may have resistance mechanisms analogous Carlson and Knight: Agrilus beetles 29 to those of conifers. For some conifers a close relationship between oleo- resin exudation pressure and tree resistance to scolytid attack has been demon- strated (see for example, Vité, 1961). While the nature of conifer oleoresin ducts and their products makes pressure measurements relatively easy, host resistance could not be studied as readily for species whose chief resistance mechanism is the exudation of phloem sap. Piercing the phloem causes an injury which interferes with its normal activity. However, the observations of Heering (1956) suggest that useful pressure measurements might be ob- tained without tapping the active phloem itself. He says that even before the heads of newly-hatched Agrilus viridis larvae have completely entered the bark, they are often repelled by exuding sap which detaches the egg shell from the bark. Without the shell to push against the larvae are helpless. If, as it appears, phloem pressure is actually the most important host resistance factor encountered by phloem feeding Agrilus it should follow that the serpentine galleries, like those often produced by g. liragus and anxius are an adaptation giving them some advantage for withstanding the copious flow of sap. In starting a zig-zag or sinuate gallery pattern, a larva makes an initial cross-grain segment, then turns abruptly and reverses its direction. When it begins to bore past the lateral extent of the gallery segment to which it is adjacent, the larva usually reverses its direction. Heering (1956) believed that the sinuate galleries produced by Agrilus vir- idis larvae are the consequence of a fixed pattern of behavior, although they do not follow a sinuate course in very decadent hosts. Barter (1965), on the other hand, found that g. liragus makes galleries with very close transverse segments in relatively vigorous hosts, while in more decadent ones the gallery becomes more sinuous resembling those of anxius. Like xylem departures, which for anxius or g. liragus increase in number with increasing host vigor, the zig-zag or sinuate gallery pattern may be elicited as a sort of avoidance reaction. There seems to be little reason for assuming that the effect of phloem sap on the larvae could be anything other than simple suffocation. Presumably, when the larvae make departures into the xylem, the plug of frass behind them protects them from the copious influx of phloem sap. The advantages which might be derived from a serpentine gallery pattern are more nebulous, however. Barter (1957) suggested that by staying beneath the previously formed parts of the gallery, the larvae are protected from the downward, translocation of sap. However, downward progressing galleries do not seem to prevail over those progressing upward in standing trees. Barter (1957) knew that galler- ies can proceed in either direction, and his hypothesis is, therefore, inade- quate, even for explaining the facts as he knew them. Moreover, there seems to be good reason for believing that movement of sap in the phloem of even the most vigorous of trees which Agrilus can infest is, at most, very slight. Trees susceptible to Agrilus infestation are not healthy and, without exception, must be subject to high internal moisture stress. According to the mass-flow theory, phloem transport occurs from areas of higher osmotic sap concentration to regions of lower concentration. During the growing season, the direction of transport would normally be from the crown toward the roots. However, when trees are under severe moisture stress, the rate of respiration in the crown may equal or exceed that of gross photosynthesis. Under such conditions there may be no transport of carbohy- drates in the phloem, and its turgidity must become significantly lower than 30 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 in trees not under moisture stress. Furthermore, serpentine or zig-zag galleries are produced even in logs, if they are infested rather soon after being cut. This is strong evidence sug- gesting that they are produced in response to local turgidity rather than to the mass flow accompanying the translocation of carbohydrates. It seems fairly apparent that even when moisture stress is too great for the translocation of carbohydrates to occur, the phloem is still sufficiently turgid for sap to in- fuse into sieve elements that have been severed. When Agrilus larvae cut through the sieve tubes, part of the exuding sap is probably absorbed by the inactive phloem and the xylem. Consequently, the pressure of sap in gallery should be lower than that in the surrounding phloem. In spite of the fact that successful infestation must invariably be attended by low turgidity in the phloem, sap should pass slowly from it until it loses its turgidity entirely or until its pressure comes into equilibrium with that in the gallery. If the foregoing premises are valid, the function of the sinuate or zig-zag gallery pattern may be unrelated to phloem transport or to minimizing the a- mount of sap that enters the gallery. It seems possible that they minimize only the detrimental effects of sap oozing from newly-severed sieve elements, as is illustrated in figure 16. When a larva turns back adjacent to the prior- formed gallery link, the phloem inside the turn is cut off between the links forming a peninsula. These peninsulas should not exude as much sap as the phloem outside the turn. The minimization of sap influx from the peninsula might make it possible for the larvae to minimize its suffocating effects by opening the spiracles on the side of the body adjacent to the peninsula. With each successive link, then, the larvae could open the spiracles on alternate Fig. 16. Sinuate gallery pattern similar to that of Agrilus anxius. The vertical arrows indicate the hypothesized direction of sap infus- ion into the gallery after severing of the phloem sieve elements (no active transport is occurring). Because the body of the larva is a- pressed very tightly to the walls of the gallery, alternately opening the spiracles on the side of the body exposed to the smallest amount of sap infusion, might allow continual respiration. Carison and Knight: Agrilus beetles 31 sides of the body. The differences in the pattern of the g. liragus and anxius galleries may be partially related to differences in the phloem structure of birches and pop- lars. It is not sound evolutionary thinking to assert that the gallery differences are attributable either to genetic differences in the insect species or purely to differences in the hosts. In other words, it is pointless to try to separate the combined phenotypic effects of environment and genotype. A. anxius or g. liragus larvae in relatively vigorous hosts construct longer galleries than those living in weaker hosts. Barter (1957) found that anxius which matured in one season in rapidly dying birches, made galleries of only 15 to 20 inches, while those in more vigorous trees matured in two years, af- ter boring 20 to 52 inches. Anderson (1944) gives similar data for g. liragus development. The reason for the vigor-related variations in gallery length are apparently unknown, but two possibilities are evident: (1) larvae feeding in relatively vigorous tissues perhaps must expend more energy in attaining the nutrients necessary for complete development, or (2) molting of late lar- val stadia may occur only under certain conditions, such as those relating to host physiology, time of year, or both. There are no published data from which it is possible to ascertain if the gallery segments for all larval stadia are longer in more vigorous hosts or if the increase in gallery length is built up during a particular portion of larval development. The Alder-Birch Borer. Very little is known about the relationship be- tween Agrilus pensus and its hosts, Alnus spp. and Betula nigra. Because it is often difficult to distinguish pensus-infested speckled alders (Alnus rug- osa) from uninfested ones, it seem likely that pensus plays a more primary role in causing the death of these shrubs than do g. liragus and anxius in in- festing their respective hosts. It is very difficult to trace out complete pensus galleries because they meander extensively through the xylem and a number of galleries may inter- sect. Consequently, no attempts were made to accumulate data on total gal- lery length. It seems likely, though, that the average length of pensus gal- leries will approach the maximum length for anxius or g. liragus galleries. Because dying speckled alders are probably not suitable hosts for pensus, these insects are always contending with a level of host resistance which is found only in the more vigorous of the hosts suitable for anxius or g. liragus. Moreover, many speckled alders show signs of unsuccessful attacks, and suc- cessful ones do not seem to occur very frequently. Much of the earlier por- tion of larval life is spent in the xylem, and the mortality rate of young larvae seems to be very high. Blanchard (1889) records one location in Massachusetts where he found pensus (under the name granulatus) in rather large numbers. Barter and Brown (1949) collected a large series of pensus on Alnus crispa var. mollis and suggest that it is particularly well-adapted to this alder, which is confined to the Atlantic seaboard. However, A. c. var. mollis does not appear to differ greatly inform or in habitat from A. rugosa. Furthermore, in June, 1968 an area in which pensus adults were moderately abundant was encountered at Mallard Creek in Iron Co., Michigan. The creek bottom was flooded by beavers during the previous summer and possibly also in the sum- mer of 1966. It, therefore, appears that, at least throughout most of its range, pensus becomes numerous only infrequently, particularly because of the host resistance controlling factor. The Aspen Root Girdler. Agrilus horni has different means for coping 32 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 with host resistance than have anxius, liragus, or pensus. First of all, rather than attacking dead or dying hosts, it limits its attacks to those poplar suckers small enough to be overwhelmed, and those on which eggs are deposited appear to be of normal vigor. The vigor of aspen suckers must be affected by the root girdling of horni larvae. Zahner and Debyle (1965) found that pruning ei- ther of the parental roots of Populus grandidentata suckers about 4 or 5 years old usually causes a significant reduction in the growth of the suckers over a span of two growing seasons following treatment. Usually, horni larvae com- pletely masticate the xylem in one of the roots in which they bore. This occurs at a point where the diameter of the root is about 1/8 to 3/16 of an inch in di- ameter, and its chief function may be to elicit the turn of the larva back toward _the stem of the host (see page 20). However, because water absorption occurs mainly in a zone 1 to 10 cm from the apex of a root (Kozlowski, 1964), the xy- lem girdling may induce moisture stress or increase it in the stem of the host and in the part of the root proximal to the girdle. If the xylem girdle occurs in a branch of a main (or parental) root, or if it sometimes does not occur at all, the survival of larvae may depend upon their ability to effectively girdle the phloem, thereby causing starvation in the distal part of the root or pre- venting food from reaching the stem and proximal part of the root from the parent tree (ortet). There are probably definite size limits on roots which can be effectively girdled in either manner. In 1960 Nord (1962) measured the diameters of 150 horni-killed suckers. He found that 84 percent of these had diameters of 0.5 inches or less. He was, however, unable to demonstrate that horni shows a preference for small suck- ers because a similar percentage of the suckers in the stand he sampled had diameters below 0.5 inches. Studying the same area in 1962, Fowler (1963) found that 71 percent of the infested suckers had diameters below 0.5 inches, while about 76 percent of the healthy ones were below that size. Using regres- sion and variance analysis, he compared the relationship of sucker height to diameter for infested suckers with that for uninfested ones, and found no dif- ference. His illustrations show the two sets of height-diameter data to be strikingly congruent. Fowler (1963) lists the numbers of infested and uninfested suckers in each of the age classes from 1 to 6 years, and his data are presented in table 2. Table 2. Age-class frequency distribution of healthy and Agrilus-killed Populus tremuloides suckers: Number of Suckers “ee Healthy horni-Killed 1 0 1 2 3 13 3 19 58 4 BOs 51 5) 93 293 6 20 4 Total 150 150 “Modified from Fowler (1963). Carlson and Knight: Agrilus beetles 33 When the two distributions are made comparable by subtracting one year from the age of the healthy suckers, it can be seen that the age-class frequencies of infested and uninfested ones are very similar. It must be remembered that suckers which died at age X were actually selected by females at age X-1. Hence, the suckers available to horni females in this area were from 1 to 4 years old. The maximum height of suckers in the area would have been about 6.5 feet. This is near the maximum size of suckers on which horni eggs have been found. The data of Nord (1962) and Fowler (1963) seem to indicate that the horni females were ovipositing randomly with respect to sucker size. It appears that when the eggs were laid, few of the suckers were too large to elicit ovi- position, or that too few were sufficiently above the optimum size for survival of larvae to make it apparent in the statistics. A small experiment was initiated in 1964 to examine the effects of host re- sistance on horni larvae. Four groups of 15 Populus tremuloides suckers were chosen with the intention of introducing first instar horni larvae at their bases. Two groups were composed of suckers beginning their second year of growth, and the other two groups of suckers were starting their fourth grow- ing season. They will be referred to as 2-year-old and 4-year-old suckers. An area was picked in which the likelihood of having the experiment con- founded by natural infestation was low, although this approach, admittedly, has undesirabilities. A random sample of the heights of 2 and 4-year old suckers was taken. Heights were measured with a leveling rod and were re- corded to within the nearest one-tenth foot of the point where the previous season’s growth terminated. The mean heights were calculated for each age class. Within each age group, height classes of +1 standard deviation from the mean were set as the sizes for suckers to be chosen for the experiment. The resultant height val- ues for 2-year suckers were 1.3 and 3.2 feet, while those for 4-year-old suckers were 3.7 and 7.4 feet. These treatments will be referred to as 2-year-, 2-year+, 4-year-, and 4-year+, respectively. On selecting the suck- ers for the experiment a +0. 2 foot leeway from the treatment value was al- lowed. Eggs for rearing the horni larvae were procured from 10 cheesecloth cages which were placed over individual aspen suckers (fig. 17 and 18). A pair of beetles was introduced into each of the cages, and the eggs were collected periodically. The eggs were removed in groups by cutting away a portion of the bark on which they had been deposited. After the eggs were removed ad- ditional soil was placed in the can which enclosed the base of the suckers so that subsequent oviposition would occur on smooth, uninjured bark. The eggs and bark to which they adhered were placed in small incubators, which consisted of 1 by 3 inch white pine blocks with five shallow depressions made with a 1/4 inch drill (fig. 19). The eggs were placed in the depressions and a microscope slide was fastened over the tops of the blocks with masking tape. To keep the incubators moist, dampened paper toweling stapled to wood strips was held in contact with the ends of the wood blocks by rubber bands. The pan in which the incubators were placed was covered to prevent dessica- tion. A camel hair brush was used for transferring larvae. They were placed in small petri dishes lined with moistened blotting paper and transferred to the test suckers as soon as possible. The larvae were placed in small cavi- ties cut in the bark about one inch above the sucker bases. The cavities were 34 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 Fig. 17. Unassembled cheesecloth cage for Agrilus horni oviposition. Fig. 18. Assembled cheesecloth cage for Agri- lus horni oviposition. Carlson and Knight: Agrilus beetles 35 Fig. 19. Wooden incubators for Agrilus eggs. Standard microscope slides (1 X 3 inches) are taped over the tops to cover the small depressions holding the eggs. prepared with a sharp scalpel by making two horizontal incisions about one eighth inch apart through the bark to the surface of the xylem. The incisions were cut obliquely downward so that when the strip of bark between them was pried out, the cavity formed would keep the wriggling larvae from falling out. After the larvae were placed in the cavities, they were covered over with tape. Test suckers were subsequently revisited for checking on the survival of the larvae. Those which had died before boring into the exposed phloem region were replaced. All dead larvae were replaced until all suckers har- bored larvae which had started into the phloem. It was necessary to replace seventeen larvae before all suckers were infested, although it was apparent when the suckers were examined in 1966 that some of the larvae probably died after boring a very short distance. Introductions were begun on July 25 and completed on July 31, 1964. A few days following completion of the introduc- tions, the cavities in the bark were filled with grafting compound to prevent fungal infection or dessication at these points. Evaluations were made by digging up the suckers and their roots on May 4 and 5, 1966. The larval galleries were traced by cutting into the stems and roots with a sharp knife. The results are tabulated in table 3. It can be seen that only one larva matured in a 2 year- sucker, and none completed develop- ment in the 4 year+ suckers. Six of the larvae which eventually formed pupal chambers were in the 2 year+ (2) and 4 year- (4) suckers. The data are, of course, rather meager, but it is likely that survival in the 2 year- and 4 year+ treatment was poor primarily because of the unfavor- able conditions for the larvae in the roots. In 6 of the 2 year- suckers the larvae had to bore in roots that were, perhaps, too small to allow successful development. Larvae are known to have successfully matured after spiraling over a span of only about 6 inches in small roots, but this is probably not al- ways possible. Survival of horni larvae was probably impossible in three of the suckers having roots of adequate size, because the roots were diseased and decomposing at various points. One of the 2 year- suckers had a very large root, about 1 1/2 inches in diameter. Such large roots probably are not suitable for horni larvae either, 36 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 Table 3. Development of Agrilus horni larvae in selected Populus tremuloides root suckers. Number of Larvae Ny a A cee Boring at Least| Boring More Completing Short Gallery | Than 10 Inches| Development ayr- 14 8 1 2yrt+ 14 8 2 4yr- 11 5) a 4yr+ 11 4 0 All 50 25 ti “Suckers were in their second (2yr) and fourth (4yr) growing sea- sons when larvae were introduced and had heights (+0. 2 ft; see p. 33) of 1.3 ft (2yr-), 3.2 ft (2yr+), 3.7 ft (4yr-), and 7.4 ft (4yr+). probably because they cannot be girdled readily. Large root size was undoubt- edly an important factor contributing to mortality in 4 year+ suckers too, but some of the larvae in these suckers seem to have perished even before reach- ing the roots. All the information so far assembled indicates that horni can kill healthy aspen suckers. However, there is an upper limit to the size of hosts it can subdue. It also seems likely that, if other factors are equivalent, the suckers on areas of good site quality are able to resist infestation better than those on poor sites. The areas in which horni is likely to establish persistent popula- tions are ones where suckers spring up in fields at the edges of stands of lar- ger aspens or in areas where the aspen clones do not expand rapidly to fill the intervening spaces. The site quality in such locations is usually low even in | the former case, but is invariably poor in the latter. A. horni is usually not found in areas of high site quality, and never becomes abundant on such sites. Nevertheless, one should not assume that their scarcity in areas of high site quality is directly related to the poor survival of larvae in these areas. The horni adults seem to orient to open areas, and good sites usually produce dense, closed stands. Moreover, the adults may have evolved this affinity for open areas at least partly through the effects of genetic selection channelled via differential survival of the larvae. Poor aspen sites which are left rather denuded after fire, like the main study area of Nord (1962) and Fowler (1963) are likely to be good horni habitats after the aspens have suckered. The numbers of horni in such places should decrease rapidly over a period of 6 to 8 years, however, Infact, in 1964, approximately four years following the year of peak abundance, there were on- ly a few horni-infested suckers to be found on the plot where Nord and Fowler had worked. The largest infested suckers listed in their data had diameters of 1.0 inches (6 inches above base). A. horni eggs were not found on suckers larger than 0.9 inches at the base. It appears as though host selection may be the factor limiting the sizes of suckers which are actually infested. Again, the crucial survival factor would appear to be root size, and this certainly corre- lates rather well with stem size. Although host resistance may not directly limit the upper size of suckers in which horni larvae can live (i.e., by simply causing the mortality of those above the upper limit) one must presume that adult host selection behavior is evolutionarily directed by the ultimate results Carlson and Knight: Agrilus beetles Je OL it, PREDATORS. Woodpeckers are the only predators known to exert any pop- ulation limiting effect on anxius and g. liragus. Their predation ong. liragus is confined to standing trees (Barter, 1965), and the same is probably true for woodpecker predation on anxius. Knight (1958) found that the feeding of wood- peckers on bark beetles has the greatest effect when the beetle infestations are heavy and infested trees are concentrated. Similarly, Barter (1957) found that their predation an anxius appears to be more prevalent in heavily infested trees. One might then expect that the effectiveness of this predation would be rather variable depending upon the size and concentration of populations, al- though other factors are probably also important. Nash et al. (1951) found that the toll of woodpeckers on anxius varied from very slight in some areas to as high as 30 percent in others. Barter (1957, 1965), however, found that woodpeckers consumed as much as 50 percent of anxius brood, and removed up to 40 percent of the g. liragus brood from their pupal cells. Anderson (1944) says that about 10 percent of anxius and g. liragus prepupae were con- sumed by woodpeckers. Woodpecker predation is confined to the spring, apparently occurring only after the larvae begin the transition to the prepupal stage. Barter (1957) be- lieved that woodpeckers fed on anxius at that time only, but later (1965) he found that the pupae of g. liragus were consumed and presumed that adults were also eaten. Woodpeckers are also an important population limiting factor for pensus. It appears that the predation on pensus occurs during the winter. Large num- bers of pensus brood are never found in a single Alnus rugosa stem, but the extent of concentration of infested alders may affect woodpecker activity. It seems that once a woodpecker finds the first pensus pupal chamber in a stem he is likely to find them all, although there usually are not more than 7 or 8 in a single stem. | : We have seen spiders and ants attacking the adults of g. liragus and anxius, but they probably cause little mortality. These beetles usually apress the legs to the thorax and fall free of the host in time to avoid being caught. In one in- stance, however, an ant had grasped its falling prey and both predator and prey fell to the ground where they couldn’t be found. The quickness with which these beetles take flight or drop to the ground, especially on hot days, is ap- parently reflected in Gory’s name anxius (nomen nudum of Dejean), and it seems rather unique that torpidus Leconte has been placed as a synonym of the former. Nord et al. (1965) observed ants mouthing the eggs of horni on caged aspen suckers. Since ants frequently pass up and down the stems of suckers, often in visits to sources of aphid honey dew, they may encounter the horni eggs which are placed above ground level. Because many eggs survive to hatch, they do not appear to be particularly attractive to ants, however. PARASITES. Nash et al. (1951) listed eight parasite species which were reared in cages containing logs infested with anxius. It is certain that some of the Ichneumonidae they list as anxius parasites were instead reared from other insects’. Glypta sp. and Pimplopterus sp. are parasites of Lepidoptera, while 1 Identifications made for Nash et al. were undoubtedly reliable as they say these were made by Townes and Muesebeck, both prominent specialists. 38 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 Olesicampe sp. is a sawfly parasite. None of these could have been reared from anxius. The parasite Barter (1957) reared from birches was identified for him as Ephialtes sp., but as more recently been identified as Dolichomitus messor perlongus s Cresson'. The Ephialtes species reared from Populus by | Populus by Barter (1965) is probably the same species. The parasite identified for Nash et al. (1951) as Ichneumon sp. is probably Dolichomitus sp., perhaps D. messor perlongus. Nomenclatural corrections in the usage of Ephialtes Schrank and Ichneumon L. have reinstated their application to groups parasitic on Lepi- doptera?. Townes and Townes (1960) record D. messor perlongus as being reared from Saperda calcarata Say, and S. concolor Leconte, and it also parasitizes Oberea shaumii Leconte. Townes? expresses doubt that females of messor perlongus could develop on Agrilus spp. The records of several braconids reared by Nash et al. (1951) from anxius are probably legitimate. The species are Atanycolus charus Riley, Spathius simillimus Ashmead, and Doryctes atripes Provancher. They also reared the chalcid, Phasgonophora sulcata Westwood. Barter (1957) reared an additional braconid from anxius, Helconidea liga- tor Say, but did not rear Doryctes. However, he records Doryctes sp. para- sitizing g. liragus (Barter, 1965). Besides the Ephialtes sp. already discussed, other parasites Barter reared from g. liragus were Atanycolus charus, Phas- gonophora sulcata sulcata, and the eulophid, Tetrastichus sp., listed as probably be- ing near rugglesi ieglesi Rowher. Barter (1957) says that Atanycolus and Phasgonophora were the most im- portant larval parasites of anxius, parasitizing 1 to 52 percent (average 14 percent) and 1 to 8 percent (average 4 percent) of the larvae, respectively. Other parasites were recovered only occassionally. He recovered parasites from only 9 percent of all larvae in his samples. Therefore it appears that total parasitism was not usually as great as is indicated by the percentages listed above, which were calculated for a sample of 600 larvae. The only parasites which was consistently reared from g. liragus larvae by Barter (1965) is Phasgonophora sulcata, which parasitized from 2 to 20 percent of the larvae. Other parasites were reared less frequently. Parasitism on larvae of anxius and g. liragus may amount to a rather in- significant portion of the total mortality factors which regulate their popula-_ tions. The same cannot be said for egg parasitism, however. Barter (1957, 1965) and Nash et al. (1951) found that parasitism was consistently high on eggs of anxius and g. liragus, usually being near 50 percent for both species. Apparently, two chalcidoids were about equally important, even though in in- i The specimens reared from Betula were lent to us by Mr. Barter for identification, but those he reared from Populus has been accidentally destroyed. 2 The revision of the Ephialtinae by Townes and Townes (1960) had led to the change for Ephialtes, and Ichneumon, sensu Townes (1944), included Dolichomitus, sensu Townes (1960). S Personal communication, 1966; H. K. Townes, American Entomological Institute, Ann Arbor, Michigan. Carlson and Knight: Agrilus beetles 39 dividual cases the species were alternately dominant. One was identified as Thysanus sp. (Thysanidae) and the other is an encyrtid near Coccidencyrtus. While the egg parasites of anxius and g. liragus must be more important in population regulation than the parasites of the larvae and pupae, the reverse seems to hold for horni. Only four eggs in a sample of 147 (about 3 percent) taken in 1965 were found to be parasitized. The parasite was identified as Oobius sp. Although they are exposed to parasites for a shorter time than the larvae of g. liragus or anxius, horni larvae are parasitized much more heavily. A. horni pupae are also parasitized rather frequently. Total parasitism of lar- vae and pupae was about 42 percent in 847 rearings, with Tetrastichus nordi Burks being reared most often. This eulophid is quite small and probably lays its eggs on the larvae when they spiral just beneath the bark in the stem, between the time of their emergence from the root and formation of the pupal chamber. This was deduced from observing the behavior of a few T. nordi females in the laboratory and from X-ray pictures of T. nordi larvae develop- ing just beneath the bark on larvae that never formed a pupal chamber. Usually, however, the horni larvae are not killed by Tetrastichus until after the pupal chamber is formed. Emergence of T. nordi adults corresponds with the time when horni larvae are re-entering the sucker stems (August and early September). An average of 49.14 17.1 adults emerged in a sample of 49 broods. The size of the adults and their sex ratio is quite variable. Three ichneumonids were found with about equal frequency in horni pupal chambers. They parasitized a combined average of approximately 13 percent of the horni collected. These parasites have longer ovipositors, enabling them to place eggs on the larvae and pupae within their pupal chambers. Xylo- phrurus agrili Viereck has been reared from horni larvae and pupae in both spring and fall. Brooks (1914) says this species has two generations per year. He found it to be an important parasite of A. vittaticollis, which bores in the roots of various trees and shrubs of the family Roseaceae. In the fall it lays its eggs on the vitaticollis larvae at about the time they are forming the pupal chamber. The spring brood attacks pupae or larvae just prior to pupation. A. vittaticollis may be the primary host for X. agrili, as Brooks found it to destroy from 25 to 40 percent of the larvae and pupae of this beetle. Xylophrurus bicolor maurus Townes emerged from horni pupal cells only during the spring, and all of the collecting and rearing records which Townes and Townes (1962) list for X. bicolor indicate that the adults fly only during the spring. X. bicolor seems to have a diapause similar to that of many Agri- lus spp., requiring an extended period of cold for development to be completed. It is perhaps for this reason that efforts to rear the adults from suckers col- lected during the initial stages of larval development are not very successful. No rearing records were previously available for X. bicolor maurus. How- ever, the western subspecies, X. bicolor bicolor has been reared from an in- sect in Salix sp. and from Saperda moesta Leconte infesting Populus balsami- fera ssp. trichocarpa (Townes and Townes, 1960). The third ichneumonid, Mastrus smithii Packard, was incapable of emerg- ing from horni pupal cells. The only adults obtained were removed from the infested suckers where they died, often after boring only part of an exit hole. potdrniined by B. D. Burks, U. S. Dept. Agr., A.R.S., Entomology Res. Div., Wash- ington, D. C. 40 | Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 The mandibular teeth of these specimens are usually worn considerably. This species is known to be a secondary parasite on the ichneumon fly Gambrus extrematus Cresson and has been reared from the coccoons of two sawfly species (Townes, 1951). It seems likely that it is poorly adapted for emerg- ing from Agrilus pupal cells, and probably parasitizes ichneumonid parasites of other insects with greater success, particularly those in the coccoons of cecropia moths.. Besides Tetrastichus nordi, several other Chalcidoidea' were reared from horni pupal chambers. They caused about 6 percent mortality of the horni reared in this study, and this was attributed principally to Lampoterma sp? | a pteromalid. This parasite emerges about mid-May, and therefore, oviposits chiefly on prepupae or pupae of horni. From 3 to 10 adults develop in each horni pupal chamber. This Lampoterma sp. appears to be limited to one generation per year by an obligatory pupal diapause. Eurytoma magdalidus Ashmead (Eurytomidae) was reared from several horni pupal cells, but this species may be a secondary parasite. In two in- stances it is known to have emerged from pupal chambers which harbored two different types of parasite larvae. Some of the larvae and pupae which died for unknown reasons (17.7 percent) may have been killed by parasite stings. Consequently, the impact of parasi- tism may be underestimated. ADULT BEHAVIOR Feeding and Maturation Barter (1957) found that Agrilus anxius females, maintained in the labora- tory, fed about 6 days before showing any tendency to lay eggs. Adults of anxius and g. liragus are not found mating or ovipositing in the field until about 2 weeks following emergence. There is no reason to suspect that the same is not true for horni and pensus. Oghushi (1963) found that under laboratory conditions Agrilus auriventris auriventris Saunders females formed ova 5 to 6 days after emergence, but full grown ova were not present until 8 to 10 days after emergence. Oviposition be- gan on or after the eleventh day following adult eclosion. Ogushi found no full- grown ovaries in females which had not fed. Their development was very slow in unfed beetles, many of which did not live long enough to reach the oviposi- tion stage. Similar developmental requirements are probably common to most, if not all, Agrilus. Some unfed anxius females dissected quite soon after emergence had partially developed ova. Without food, however, these females are prob- ably unable to form fully-developed eggs, and usually die after a rather short time. A. anxius adults were maintained successfully during the winter on pieces Ata nne by B. D. Burks, U. S. Dept. Agr., A.R.S., Entomology Res. Div., Wash- ington, D. C. Nord et al. (1965) list this species as Metastenus sp., but the application of Metastenus has recently been changed (cf Burks, 1967). Carlson and Knight: Agrilus beetles 41 of apple, but in nature their feeding is, no doubt, restricted to leaves. Hutch- ings (1923a) says the bronze birch borer feeds more readily on the foliage of poplars and willows than on birch. Barter (1957) found that anxius adults con- sume the leaves of poplars, particularly those of Populus tremuloides, more readily than those of Betula papyrifera. The findings of Nash et al. (1951) differ slightly; in their laboratory studies anxius showed the greatest prefer- ence for P. grandidentata foliage. They rate the species used in their tests in the following order of decreasing preference: P. grandidentata, P. tremu- loides, Salix spp., Betula lutea (= alleghaniensis), B. papyrifera, P. balsam- ifera, and Alnus spp. Very little is known about the feeding habits of anxius adults, but they were occasionally observed feeding on B. papyrifera leaves by Barter (1957). Brit- ton (1922) says bronze birch borer adults (those attacking birches) fly about and feed to some extent on the foliage of willow, poplar, and probably birch. We have not tried feeding horni adults anything besides P. tremuloides leaves in the laboratory, and horni was found in the field on no plant other than tremuloides. Aspen leaves undoubtedly comprise the major portion of the adult horni diet, which is probably true for g. liragus also. A. pensus, on the other hand, did not readily consume the foliage of as- pens. Because they have been collected on the foliage of alders, but not on other plants, one would expect them to show some specificity for alder leaves. However, the only foliage which they seemed to eat readily in the laboratory was that of Salix spp. Very little testing with alder was possible, however, as the beetles were maintained in Ann Arbor, where alders are not readily available. Usually, pensus adults did not survive for more than a few days, no matter what kind of foliage they were provided. Adults of other species kept in the same controlled environment chambers survived exceptionally well (at 70 percent relative humidity; day temp 75°, night temp 60° F). The poor Survival of pensus adults may have resulted because of a failure to provide the proper temperature and humidity for pupal development. Dispersal Nothing has been written about the dispersal capacities of g. liragus or anxius, and little is known about the dispersal of any Agrilus, including the ones that are naturalized pests in the United States. Heering (1956) has ap- parently made the only attempts at release and recapture studies with marked Agrilus. However, as only 200 A. viridis adults were released he was unable to recapture enought to give a reliable result. Although Agrilus beetles may be capable of flying considerable distances, they probably do so only rarely. In rearing and maintaining these beetles in the laboratory we have seen nothing which would indicate the existence of any sort of antagonistic mechanism by which ‘‘flight exercise’’ acts as a releaser for feeding or reproductive activity as has been demonstrated for some aphids (Kennedy and Booth, 1963) and scolytid beetles (Graham, 1959, 1961). If they flew only far enough to encounter suitable hosts, they would usually be staying quite near the places where they emerged. Host Selection Following the period of reproductive maturation the adults are attracted 42 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 to the hosts which have the particular attributes making them suitable for mat- ing and oviposition. Many Agrilus and other Buprestidae seem to show a strong tendency to select hosts which are well exposed to sunlight. Consequently, those which live on trees are most active at forest edges or in park-like stands. ANXIUS AND G. LIRAGUS . Generally, anxius females are said to oviposit on the sunny sides of trees and on those most exposed to sunlight. However, Bar- ter (1957) found that the photopositive tendencies of anxius can be overridden, to some extent, by other stimuli, presumably olfactory ones. Barter says that when the shady sides of trees are wounded, females oviposit there rather than on the unshaded side. He concluded that the olfactory response is often strong- er than the photopositive response. Anderson (1944) came to similar conclusions. During the first season of his two-year study he found no deviation from the commonly accepted idea that trees exposed to full sunlight attract significantly greater numbers of anxius and g. liragus than shaded ones. Using a 30 second counting interval Ander- son found an average of 2.6 beetles per unshaded tree, while only 1.8 were found on shaded trees. When the experiment was repeated the following year, however, he found that the count for unshaded trees did not differ significantly from that for shaded trees. During the first season, trees with the tops bro- ken off by winter damage were abundant, but such damage did not occur during the second year. Anderson thinks that greater numbers of weakened hosts were available during the first season, allowing g. liragus to make a selection for those more exposed to sunlight. His conclusion is that the physiological condition of the host is more important than its degree of exposure to sunlight. There may be sexual differences in the photopositive response, but little has been done so far to account for the possibility. Males and virgin females may be more photopositive than ovipositing females. Females ovipositing on fallen trees are apt to work along the sides of stems on days when males orient more towards the upper surface. One is prone to confuse the effects of light and temperature. It seems quite likely that these beetles will appear to be more photopositive on relatively cool days (but still above 70° F) than on hot days. An observation made in 1964, during studies of anxius behavior, illustrates this point. In order to concen- trate the beetles on bolts cut from freshly-felled birches, the tops and other unused parts of the birches were pulled well into the shade. This slash, never- theless, attracted some of the beetles, and the bolts under observation, which were in full sunlight, attracted only moderate numbers of them. That day the temperature maximum was 85° F (U.S. Weather Bureau, 1964; Stambaugh, 15 July) and the sky was clear, so that temperatures may actually have been too high for the insects to remain for extended periods in the direct sunlight. In another of Anderson’s (1944) experiments, the attractiveness of 7 types of host material was evaluated by tallying the numbers of adult g. liragus on each aspen test tree during a 30 second counting interval. Observations were made between 10 AM and 3 PM on sunny days when the shade temperature was between 75° and 90° F. His results are presented in table 4, and it is quite evident that injured trees are more likely to be attacked than uninjured ones. Barter (1957) found this to be true for anxius also. From Anderson’s data (table 4) one may also surmise that standing trees are more attractive to g. liragus than similarly treated trees which are felled. Both the topped trees and topped and girdled trees attracted significantly greater numbers of beetles than trees which were felled. Perhaps this occurred because the standing trees Carlson and Knight: Agrilus beetles 43 Table 4. Attractiveness of various types of Populus tremuloides host material to Agrilus granulatus liragus. Condition of Average Number of Beetles | Number of Host Trees per Tree per Observatior “| Observations 1. Vigorous 0 2. Suppressed 0 21 3. Girdled 9 sl. 54 4. Topped | Lied 54 5. Topped and girdled 2.8 27 6. Felled and lopped 1.0 7. Felled, not lopped £3 “Modified from Anderson (1944). : ’The values are corrected so that each represents the number of bee- tles observed per 17 ft of bark area, the average for all test trees. “The only statistically significant differences are those between num- bers not included within the span of a single vertical line. were more fully exposed to sunlight than felled ones. In any case, these re- sults could be misleading because standing trees rarely are as decisively in- jured by natural factors as are trees which are artificially topped or topped and girdled. The only natural injury which is really comparable occurs when trees are blown over. Less decisive natural injuries occur to standing trees when the tops are broken by winter ice damage or when the tops of already decrepit trees break where the wood exposed by repeated insect attack and disease be- comes dry and brittle. Aspens are frequently victims of this type of injury, since the wood of these species is not particularly tough, and because they are often severely attacked by wood borers like Saperda calcarata and fungal para- sites like Hypoxylon pruinatum for a period of years before they finally succumb. One might also conclude from the data of table 4 that both suppressed and vigorous trees are unattractive to g. liragus. In testing this more thoroughly, Anderson used an extended observation period of 14 days, to detect occasional beetles that might be attracted to the trees. This was facilitated by placing sheets of fly paper on the trunks of 10 paired aspens, each pair composed of a vigorous and a suppressed tree. During the 14 day observation period 18 beetles were caught by the fly paper on suppressed trees, but none were caught on vigorous trees. It is apparent that g. liragus is able to detect trees which are in poor condition, whether or not they have been wounded. The stimulus is undoubtedly olfactory, and is presumed to arise from the decomposition of sub- stances in the bark or phloem. The attractiveness of the host may, in turn, depend upon the rate of decomposition, which is no doubt enhanced by injury. HORNI. Theoretically, the kind of host selection exhibited by anxius and g. liragus should allow adult feeding to be separated from the activities involved in host selection. If this is true, then there might not be any highly developed specificity shown by feeding adults. In the late 1800’s, buprestid collectors clearly recognized that the identity of an Agrilus larval host was not necessarily ag Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 indicated by the kind of foliage the adults are taken on. Perhaps this point is over-emphasized, because the adults of some kinds of species, like horni, may show more specificity in feeding, because feeding may be involved in the host selection process. Agrilus horni does not fly directly to the stems of as- pen suckers, but instead lands on the foliage. The adults of this species usu- ally feed before moving onto the stem where mating and oviposition occur. Whether or not they always feed before crawling from the foliage to the stem is not known. Nord et al. (1965) found that after caged horni females oviposit they move up to the foliage of suckers, resting and feeding for about 10 minutes before decending to lay another egg. Under natural conditions the females may usually ascend the stem following oviposition and fly to another sucker before feeding. A. horni can maneuver with agility in flight, and can hover before landing. Their flight behavior indicates that they probably recognize aspen foliage be- fore landing on it. If so, olfaction is clearly part of the recognition process. If the odors which direct them to the host do not, at least in part, emanate from the foliage, one would expect them to land frequently on the foliage of plants other than Populus spp. This was observed only once, when a male which had just mated flew from P. tremuloides sucker to a fern next to it, staying briefly before he flew back to the same aspen sucker and began feeding. To go no farther than suggesting that horni recognizes its host by some characteristics of the foliage would be forming a hypothesis of very little value. Obviously, other factors must strongly influence the host selection of these beetles, as they do not live on just any species of Populus, or just any P. tremuloides sucker. | Previous Studies. Fowler (1963) studied the relationship of height, age, and stand density of aspen suckers to infestation by horni. He counted the num- ber of infested and uninfested suckers on systematically located millacre quad- rats. A graph of Fowler’s data (fig. 20) appears to indicate that the unit-area density of horni-infested suckers is rather directly related to the unit-area density of all suckers. However, it is not necessarily true that unit-area densities per se of the aspen suckers were the important variables affecting the infestation levels throughout Fowler’s study area. It seems likely that some of the lower stand density values can be attributed to parts of it where the ini- tial establishment of suckers was poor because of severe competition with other plants which dominated. Besides patches of sweet fern (Myrica asplen- ifolia [=Comptonia peregrina]), leather leaf (Chamaedaphne calycultata), and blue- berry (Vaccinium spp.), there were young red pines (Pinus resinosa) scattered about. Quadrats with high sucker counts, on the other hand, may simply have occurred in places where some of the suckers were not shaded by other plants. In fact, high-count quadrats probably fell where suckers were clumped (fig. 21), and sucker clumps probably had originated in spots that were left denuded af- ter the 1957 fire. Fowler (1963) notes that when infested suckers were associ- ated with clumps, they were usually at the outer edges of them. A majority of the infested suckers in this may have occurred at the outer edges of clumps. Therefore, the contagion evident in the distribution of infested suckers may have resulted from strong contagion in the distribution of suckers suitable for Oviposition. _ much more detailed analysis of the data is presented by Carlson (1968). Carlson and Knight: Agrilus beetles 45 Infested Suckers per Millacre Quadrat aR ae a a ia ie Ce LE age oe ae? ae Total Suckers per Millacre Quadrat Fig. 20. Unit area density of horni-infested Populus trem- uloides suckers as a function of stand density of all tremuloid- es suckers in the area sampled by Fowler (1963). New Studies Methods. The possibility that some contagion in egg distribution could occur independently of the distribution of suitable hosts has not been over- looked. In the spring of 1964 an area with an evenaged stand of aspen suckers was selected for studying this problem. It is an area near Beechwood, Michi- gan, where furrowing for the planting of pines had caused prolific root sucker- ing by scattered, mature Populus tremuloides, some of which were still stand- ing, while others had been pushed over (fig. 22). Because the problem involved determining the spatial distribution of in- fested or egg-bearing suckers in relation to each other, there was little need for using standard sampling methods for estimating density (i.e. stems/unit area). Instead, it was decided that the problem could be best approached by using distance measurements as indicators of spatial relationships. The two variables measured were: (1) the distance from predetermined sampling points to the nearest infested or egg-bearing sucker (DNS) and (2) distance between each infested or egg-bearing sucker and the nearest neighboring infested or egg-bearing one (DNN). Sampling points were located systematically. Five plot centers were established, four at the corners of a 400 foot square and one in its center and also approximately centered within the study area. Around each of the plot centers, 8 sampling points were established ona _ circle with a 50 foot radius. Sampling point number 1 was always located due north of the plot center, and the others were located at subsequent azimuthal intervals of 45 degrees. Sample distances were measured with a 100 foot tape. 46 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 Fig. 21. A spot in the study area of Nord (1962) and Fowler (1963), showing the clumped distribution of Populus tremuloides suckers and abundance of com- peting vegetation (from Nord, 1962). The aspen suckers occurred under similar conditions throughout much of the area. Fig. 22. Agrilus horni study area near Beechwood, Michigan. The stand of Populus tremuloides suckers, which is relatively uniform in distribution,devel- oped following furrowing for the planting of pines. Equipment used in the furr- owing operation pushed over many of the mature aspens. Carlson and Knight: Agrilus beetles 47 The Wilcoxon matched-pairs signed-ranks test (Siegel, 1956) was used for ana- lyzing the data. Discussion of the results is deferred until later, where the results of a number of analyses are discussed together. While evenaged sucker stands are best for studying the distribution of eggs by females, independent of sucker distribution, they are not well suited to studying the sucker size selection of hornifemales. For this, unevenaged areas are more appropriate. Unevenaged sucker areas occur in the ecotones between aspen forests and open fields or in park-like stands where clones or multiclone groups are apt to stand isolated, annually producing some suckers at their edges. Such areas are also of particular interest because they har- bor more perpetual infestations than do evenaged stands. Devising sampling techniques for unevenaged areas was problematical, however. The most per- plexing difficulty is that in these areas the zone in which the horni infestation occurs represents a rather small proportion of the ecotone area. Hence, it is difficult, if not impossible, to devise a random sampling scheme that is efficient. Even a systematic approach is difficult to contrive because the habi- tat of interest is usually distributed in a rather irregular manner. Therefore, the usual kinds of systematic procedures are not applicable. The compromise devised cannot be considered ideally objective, but that does not seem to greatly impair its value. The sampling was carried out by proceeding along the edges of the clumps or closed stands of larger aspens and selecting the first 6 ft sucker encountered as a plot center. In turn, the, Tit, 5 ft, 4 ft, 3 ft, 2 ft, and 1 ft suckers nearest the 6 ft one were selected. The distance between the 6 ft sucker and each of the others (D,) was meas- ured. Before the suckers were chopped free at their roots and examined for eggs, the distance to the nearest neighboring sucker in the same height cate- gory (DNN) was measured and recorded. Only the first set of suckers selected was examined for horni eggs. Besides tallying the numbers of eggs and noting their positions, we also recorded the basal diameters of the suckers (DAB). Successive plots were always separated by at least 3 times the maximum D,. Six foot suckers were chosen for the sampling centers because they were near the upper size limit for suckers which would bear horni eggs. Conse- quently, choosing a 6 ft sucker would insure that sampling would include areas where suckers above the upper size limit occurred, while still keeping it with- in areas that were likely to be horni habitat. This sampling method was employed in one area near Kenton, Michigan and in another near Phelps, Wisconsin. Analyses of the data chiefly involved the use of chi-square comparisons and graphic procedures. Expected values (E,) for the chi-square analyses were derived from expectancy factors (E,), which are weighting factors based upon D, and DNN (collectively, D values) measurements. The basis for the Seen ent ion of expected values is the as- sumption that the abundance of suckers ina particular size class is inversely related to the area of a circle which has a radius equal to the average D value for the class. Hence, density is proportional to 1/ D7. In deriving the E, val- ues it was anticipated that DNN measurements should account for any clumping tendency among suckers of the same age. On the other hand, the D, measure- ments should account for any stratification that existed, such as gradients in the positions of suckers relative ,t0 those of other sizes. E, values are com- puted simply by dividing the 1/ D? value for each size class be the total for all 1 Each height class had a+0.5 ft interval. 48 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 classes (giving each E, relativity to all others). The expected numbers of egg- bearing suckers are derived by multiplying the total observed number of egg- bearing suckers by each E, value. One set of chi-square comparisons was made for height classes, and in another set the numbers of egg-bearing suckers were apportioned into DAB classes (table 6). Diameter class (DAB) values for the latter comparisons were computed from regressions of DAB over sucker height by substituting the values for the height class limits into the equation and solving for DAB. To compare height and DAB as indicators of the suitability of suckers for horni oviposition, ratios of observed to expected values are used. This re- quired the computation of additional expected values for a set of DAB classes with uniform limits. It was necessary to derive expected values for these uniform DAB classes indirectly, because there was no direct (significant) sta- tistical relationship between DAB and D values. In order that all ratios of observed to expected values have positive signs, it is necessary to adjust all expected values (E,) upward so that none is ex- ceeded by the observed (O,) value for the same class. What was done, in effect, was to multiply all E, values by the greatest O,/E, ratio occurring among all sucker classes (yielding E,.). The subsequently derived O, /B,: ratios can be used for making between-area data comparisons as well as for the height-DAB comparison mentioned above. Factors with similar uses are derived by dividing the total number of eggs (S) occurring on the suckers of each size class by the corresponding E,, values. Discussion. There were no significant differences between DNS (dis- tance to nearest sucker) and DNN (distance to nearest neighbor) measurements for either the infested or the egg-bearing suckers at the Beechwood, Michigan evenaged sucker stand. Therefore, there is no marked tendency toward con- tagion in the distribution of either infested or egg-bearing suckers. Two ob- vious explanations are possible. (1) A. horni females may make a post-ovi- position flight, before they re-alight and lay another egg. This sort of behavior should lead to a rather random distribution of eggs. (2) On the other hand, suckers which were suitable for oviposition and subsequent infestation may have been rather randomly distributed throughout the area. The latter explanation seems to be the most logical. The few direct field observations made on horni females would indicate that they really do not fly very far before landing on another sucker. Nor is there any indication that a post-ovipoisition flight is an unvarying occurrence in a fixed behaviorial pat- tern. Nord et al. (1965) found that after ovipositing, caged horni females im- mediately crawled up the sucker stem and out onto a leaf. After spending about 10 minutes feeding and ‘‘resting,’’ they might descend to lay another egg, and SO on. : Under natural conditions horni females may usually fly to another sucker after each egg is laid. It may be uncommon for them to feed on a sucker just after ovipositing on it, particularly if feeding is an integral part of the host selection process. In the Beechwood area only a few suckers were found with two horni eggs, and only one had three. When more than one egg had been placed on a given sucker, there was usually no way of knowing from their ap- pearance if they had been deposited at about the same time or at different times. Unlike the Beechwood area, the egg-bearing suckers in the Phelps and Kenton areas usually had more than one egg (average 2.5) and one had a surprising total of 15 (table 5). When the difference between the latter areas and the Beechwood area are considered, it is only reasonable to conclude that the oc- Carlson and Knight: Agrilus beetles 49 Table 5. The occurrence of Agrilus horni eggs in the Kenton and Phelps areas on Populus tremuloides root suckers in height classes of 1 to 7 ft. Number of Eggs Average per Suckers Bearing Eggs Class (feet) Total | Total | Egg-Bearing Sucker 1 1 1.0 2 12 Lea 3 41 4.5 4 30 ag | 5 28 ve 6 14 Lye 7 8 LA All oe Os currence of more than one egg on a single sucker is usually not an indication that they were laid in sequence by the same female. Instead, what seems to be indicated is that some suckers were far more suitable for horni oviposition than others, even of the same size. Moreover, in the Phelps and Kenton areas the horni habitat was restricted to a very narrow zone, and movement along it probably resulted in these beetles contacting suckers in particular locations more frequently than others. The most reasonable conclusion seems to be that when suckers in these particular locations were otherwise favorable for oviposition there were a number of eggs laid on them. Sucker size should be an important factor affecting the distribution of eggs by horni females and/or the survival of the larvae. In both height and DAB chi-square comparisons the difference between the observed and expected dis- tribution of eggs is highly significant (table 6). The relationship of egg occur- rence to DAB is more sensitive than that with sucker height (figs. 23-26). The Kenton trend line of O,/E,, over DAB (fig. 24) shows a double peak, but the second peak is reduced in the graph of S/E,, over DAB (fig. 26). This suggests that in the Phelps area the suckers most likely to elicit horni ovi- position had somewhat larger diameters than those in the Kenton area. This appears to be true despite the fact that the abundance of small suckers rela- tive to those of larger sizes was greater in the Phelps area. Examination of the data on egg placement for the two areas yields the most likely reason for this difference. In the Phelps area more eggs were located in association with moss than on other kinds of deposition sites. Most of those Phelps area eggs deposited in association with moss occurred on the 0.6 and 0.7 inch suckers (all but 2 out of 21), and these two DAB classes were the source of the great- est difference between the areas. Therefore, it seems very likely that the occurrence of mosses on these suckers in the Phelps area is responsible for the positive skewing of the trend lines (figs. 24, 26) as compared to the more negatively skewed ones for the Kenton area. Inasmuch as factors independent of sucker size, such as the occurrence of mosses, affect horni oviposition, even two areas with nearly identical arrays of sucker sizes will not necessarily have similar frequency distributions for those bearing eggs. In general, however, the data do strongly indicate this much: suckers between 0.3 and 0.9 inches DAB are most likely to elicit ovi- position. ‘Jaaoy AjITIGQeqord [00 94} 4B JUBOTFTUSIS AT[VOTISTIEIS * > © ‘ZUISSIUL O18 SONTVA {YC OA} osnVdeq UOSTIedUIOD YSIoY OJ yey} WOTf SAFJIP [VIOL | rf & 489 ° 09 00 ‘FZ bz een o Z9°ST LY ‘0 € 6IL‘°0< L r e6 *eT LL‘0 7 6TL°O - Z19°0 9 = OFS 92 “T v TT9°0 - POS "0 G = 8S ‘OT (1) ae 8 €0S°0 - 96€ °0 v > OT'0 6S°E € G6E°0 - 882 °0 e 66°S 0g °9 “4 L8Z°0 - O81 ‘0 Z Q 0S ‘IT 0S ‘TT 0 I sdjeud % axf0 CT. 00°82 8Z Eco mem = eL°0 €L°0 0 i L A €% 0 Zert z 6S6°0 - 608°0 ¢ 9 H 61°9 97 °Z 9 808°0 - §S9°0 v G = LS ‘0 LG" G ZS9°0 - 90G°0 6 7 < vS'P 91 °S OT G0S‘°0 - 6SE ‘0 G g 2 G12 €8'9 € 8S$‘0 - 222 ‘0 p rd ee 29°F SL's rd IIZ"0 > I I uojUe yy g ee &. Cogs Ss) (“O) (soyout) Coz xta), « no indication that these Agrilus sounds are species-specific. These beetles probably do not produce other kinds of sounds with this same stridulatory ap- paratus. As previously noted, stridulation has not been observed in a situation that would definitely indicate that it was not stress-induced. There were a few indications that it may elicit repulsion of individuals that contact stridulating beetles. Alexander, Moore, and Woodruff(1963) suggest that, inasmuch as the sounds of Coleoptera are usually too soft to function at long distances, they may function in several fashions among adults in close proximity. The stridu- Fig. 41. Postero-ventral view of the head of an Agrilus horni °, showing the genal ridge (X109). Note the fine striations which are barely visible at one point near the posterior end of the ridge; they may be important in stridulation, but the sharp posterior portion of the ridge’s outer edge may also contact the prosternal lobe (fig. 42). Ane, G. B. Vogt, U.S. Dept. Agr. , Ac R.S., Entomology Research Division, Washing- ton, D.C. believes that ‘‘genal ridge’’ is the best possible term for this structure (per - sonal communication). Carlson and Knight: Agrilus beetles 71 Fig. 42. Finely imbricate interior surface (head is excised) of the right half of the prosternal lobe of an Agrilus horni ? (X82). latory apparatus of the species which were examined thoroughly a 2. liragus, pensus, and horni) was not sexually dimorphic, and recordings indi- cate that there is no sexual difference in the sounds produced. They do not appear to be directly associated with courtship. Although unreceptive females may occasionally stridulate, their stridulation does not seem to have any ef- fect upon courting males. In homosexual encounters, a sexually aggressive male is usually repulsed only after he mounts and probes the other, and stri- dulation does not seem to be involved. Nevertheless, stridulation may function in some other situations to prevent over-congregation of males or females, or they may serve to repel individuals of unrelated Agrilus species which may live on the same host. These stridulations should evolve toward nonspecificity in the eliciting stimulus (Alexander, et al., 1963). It is, therefore, difficult, if not downright impossible, to assess their significance by studying them un- der artificial conditions. On the other hand, as these Agrilus stridulations cannot be heard, except through amplification or by placing the insects next to the ear, and because the movements cannot be seen at distances of more than a foot or So, anyone endeavoring to determine their real function under natural conditions, will, to say the least, be faced with a real challenge. EVOLUTION Phylogeny When our work on Agrilus was initiated, we had not ils ae seriously e- nough the possibility that anxius, pensus, horni, and g. liragus are not all closely related. It seemed that the important q questions to be answered con- cerned isolating mechanisms and clues to how these species evolved from a hypothesized common ancestor. Because of this improper assessment, more species than originally anticipated will be brought into discussions about phylo- geny. THEORETIC BASES. It is quite usual to regard some characters as more reliable than others as indicators of phyletic relationships. Nevertheless, we Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 12 *poods IOJOUL MOT pues ‘1vds8 YSIy ‘yeTZ odvys ‘YIM pueq OpIM ‘Q'S UIVS [OAD] YIVWI :OTOM SSUI}OS TOZATIAQGIA oy, ‘eTqrssod SB Yep SB SSUL{IVW oY} OYBUL OJ posn 919M SUNI OM], ‘“SUIPLOOOL [VUTSIIO OY} Jo poods ode4 ey} F/T ye ‘eAoqe pouoTjUoW Ieytjdure oy} Aq Ul pes sTeusis yyM ‘azezATIAqiA Aueduiog o1sjOo Ty ACY B YIM opewM sem YdersorjOedsoIpne oy, *“1ep100 -~91 94} JO UOTyeIOdO Y}IM Po}BIOOSS¥ SpUNOS WLOI] DOUBIOJAIJUI SUIZIWITUIW ‘1apIODeI 9Y} WOIj VOUBISIP B ye pooV{d oq 0} DUOYMOIOIW OY} POMOTI[V STUT, “Tep10001 oY} poyeredo OYM ‘1910 9} 0} STSUSIS OAVS puv DUOYdOAOIW OY} ABA SoeTJOeq 94} Pley euO - ‘SSUIP1OOEI BSULYVU IOJ portnbeit 919M suOsaod OM], ‘“‘puodes red SaYyoUT ct Jo poods odey & 48 ‘ouoydoro Tu V-€ed [OPOW UBoILOULY Ue pUe ‘LOTJI[dWIe ~P-E9Ld [EPO PAODOUSeYI B ‘TAPIODIAL ZHZVEILd [OPOW P1lOoVOUsSeY, B YIM OpeUl SBA SUIPIODE.I ode} OUT, ‘9 SNSBIT] SMB[NUCIS SN[lIsV ue Jo spunos AiOVe[NpPIAIs Jo yderSor1jJOodsoIpny “EP “SIA Carlson and Knight: Agrilus beetles 73 shall set forth the assumptions concerning those we use, and give our reasons for not considering others to have the value accorded them by other students. Let it be made clear at the onset, that we do not regard any characters as being ‘‘nonadaptive.’’ Many systematists have used the term nonadaptive in reference to stabilized characters, while referring to those subject to much variation as ‘‘adaptive.’’ The irrationality of the idea of adaptivity and nonadap- tive is discussed by Dobzhansky (1956), who maintains that all traits, or at least the vast majority of them, are adaptive or have been adaptive at some time in the history of their development. To be sure, some traits are more stable than others, but their stability should relate to the kinds of functions ra- ther than to the existence of functions versus the lack of them. In many cases it is difficult to perceive functions for some seemingly trivial traits, but this is a poor reason for assuming they do not exist. So it is with the pygidial spine of species in the Agrilus anxius group. Without searching for its function, one might take what seems to be the prevailing attitude in such cases--that its stability indicates that there is no selection against it, probably because it is of no great disadvantage to its possessor. Quite to the contrary, it seems that the stability of the pygidial spine stems from its importance in the process of sexual recognition. Any large changes in the form or size of the pygidial spine calls for corresponding changes in the form of the male aedea- gus and other structures. In genetic terms, it probably owes its stability to an association with pleiotropic genes whose total influence on the developmental system is spared extensive changes through the moderating influences of homeo- stasis or coadaptation (see Dobzhansky, 1956). The pygidial spine probably has at least as much stability as characters ordinarily given tribal or generic value in other insect groups. Perhaps Oben- berger was the one taxonomist who should have had sufficient worldwide famil- iarity with Agrilus to recognize this, but he was convinced of the impossibility of even dividing Agrilus into acceptable subgenera (see Obenberger, 1959). What Obenberger (1957) implies is that, to be valid, each proposed subgenus must have as its basis a single character which occurs among no species of other groups. If any Agrilus character could meet this absurd requirement, it would probably be the pygidial spine. Obenberger (1957) denies this possibility because (1) a pygidial spine is found in the Palearctic forms Agrilus guerini Lacordaire, A. fleischeri Obenberger, and A. ater L., but is absent in an al- leged intimate ally, A. suensoni Obenberger; (2) its presence among about 30 African species makes it self-evident that it occurs among entirely unrelated groups. To counter the first point one need only ask what criteria he uses to ally suensoni with ater and the rest. He offers none, and almost any should be more subject to convergence than the pygidial spine is to total disappearance. Oben- berger’s second point is even more indefensible because the presence of the spine among African species cannot, in itself, demonstrate that it has arisen repeatedly. There is no 4 priori reason for believing that even the Ethiopian species he mentions do not share common ancestry with Palearctic ones. After all, the Palearctic region does extend into northern Africa. INTERPRETATION. Fisher (1928) has given the form of the tarsal claws pre- cedence over the pygidial spine in his classification. Asa result, three spec- ies which are here placed in the anxius group are classified quite apart from it by Fisher. These are Agrilus ruficollis, A. champlaini Frost, and A. browni (= lateralis, sensu Fisher [1928]; see p. 6). The bifid claws of these species 74 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 have the inner lobes bent toward each other. This trait appears to be the only one which they have in common with the species Fisher places them near. The pygidial spine, the marginal sclerite of the hind segment, and the general form of these insects links them with species in the anxius group. On the other hand, ruficollis, browni, and champlaini have more in common than just similar tarsal claws. They resemble each other in the structure of the pronotum, elytra, and male or female genetalia. In addition, ruficollis and browni are similar in coloration. Their hosts are not closely related, but the larval habits of all three species are somewhat alike. The larvae of ruficollis bore in the stems of brambles (Rubus spp.). During the early stages of devel- opment they often (cf Smith, 1892) form galls by making a tight spiral in the phloem, while late-instar larvae move rather directly up and down the pith. A. champlaini forms galls on ironwood (Ostrya virginia) and blue beech (Carpinus caroliniana), but the late-instar larvae spiral mostly in the wood within the gall which forms as the result of the phloem blockage caused by the gallery of the early instars. Fisher (1922) reared browni from bayberry (Myrica pensylvan- ica), and Knull (1922, 1950) says Myrica asplenifolia is also a host. The browni larvae make no galls. According to Fisher (1922), after passing the winter in the roots, the browni larvae spiral upward in the limbs for a considerable dis- tance before they reach maturity. The number and variety of similarities between ruficollis, browni, andcham- plaini makes it unlikely that they could all be convergent. However, these spec- ies differ in ways that seem to push their divergence to a rather remote era. For example, it does not seem likely that a species living on Ostrya and Carp- inus could be intimately allied with species whose hosts are Rubus spp. or Myr- ica spp. Consequently, in figure 44, champlaini is shown as also having remote connections with species that live on Fagaceae, which includes Agrilus bilinea- tus, A. acutipennis Mannerheim, and a number of other forms. While cham- plaini might instead be related to the line represented by anxius and its allies, it appears to resemble bilineatus and related species in a greater number of minor respects. For instance, the latter have the inner lobes of the tarsal claws turned inward, although to a lesser extent than does champlaini. Allies of horni. A. browni seems to be more closely related to horni than to ruficollis (fig. 44). While its tarsal claws are like those of ruficollis, browni resembles horni in having the vertex less strongly impressed, the front flattened, and the marginal sclerites of the abdomen clothed with white pubescence. The browni aedeagus is intermediate in form to those of ruficollis and horni, but browni has a trace of the prehumeral carinae, which are completely lacking in ruficollis. While the eggs of ruficollis are placed at the axils of the branches (Hutch- ings, 1922b) or leaves (Smith, 1891) of its hosts, those of browni and horni are placed at the bases of the host plants. The larval habits of browni and horni are very similar (excluding host differences). In each case the larvae bore into the roots and over-winter there. The only difference is that horni bores farther out into the root and spirals back to the stem, while browni spirals mostly in the branches. The early-instar ruficollis larvae, on the other hand, over-winter in the stem and probably spiral only during the time prior to this initial over- wintering period. Agrilus vittaticollis has the vertex strongly impressed as in ruficollis. Fisher (1928) says vittaticollis is allied with benjamini Fisher and audax Horn. The host of benjamini is unknown, but audax infests the upper branches of Ulmus ‘sotoeds Sulyew-|[[es sopnjout orty 1D an “ure}19e0uN ST AWoTjIoeds jsoy poyeusisep oy} YOTYM TOT sotoods OSOY} JO SOUBU OY} SMOTIOJ Yew uOTsSonb vy YST194Se UB YIM poyTeUl ore susuITOeds OU Uses OABY STOYINE oY} YOTYA Jo Vxey jo soweu oyy, ‘*dnor3 SniKe STI -T18V 9Y} UI pooe{d sulzoy onoaRoTeg pue OTJOLBON PEqILOSoep oY} JO oUOS AOJ 90.14 OYeTAYd [eoneu.OdAY a “Sly IO oe Tlewg jo sze10g| Carlson and Knight: Agrilus beetles eee juriceki * fleischeri fleischeri * fleischeri kurosawai * opuli satoi (?) * anulatus liragus semiaurovittatus (?) * pranulatus granulatus bilineatus criddlei tamanukii * acutipennis olivaceoniger & other forms ruficollis niveiventris fulminans ater ater vittaticollis uadriguttatus amawakii (?) * tscherepanovi * ater subcuneiformis (?) * ranulatus | fleischeri coreicus * fleischeri nipponicola * 76 Contrib. Amer. Ent. Inst., vol. 4, no. 3, 1969 rubra (Knull, 1934).