Control Unit, Grand Cayman, Cayman Islands, West Indies; David J. Lewis of the Medical Research Council, British Museum (Natural History), London, England; John R. Linley, Entomological Research Center, Florida Department of Health, Vero Beach; J. MaldonadoCapriles, University of Puerto Rico, Mayaguez; Glenn M. Stokes of New Orleans, La.; Roger W. Williams, Columbia University School of Public Health, New York; and the staff of the American Museum of Natural History (AMNH) in New York. Holdings at this museum include extensive material from the Bahamas, mainly from the Van Voast-AMNH Bahamas Expedition of 1953.

Our West Indian Culicoides collections, unless otherwise specified, have been deposited in the U.S. National Museum, Washington, D.C. Whenever possible we have deposited duplicate material in the collections of the British Museum (Natural History) in London, Institute of Jamaica in Kingston, University of Puerto Rico in Mayaguez and the

School of Public Health in San Juan, American Museum of Natural History in New York, University of Florida and Florida State Collection of Arthropods in Gainesville, and the University of the West Indies in St. Augustine, Trinidad.

We have adapted many of the illustrations from our earlier publications. They have been drawn by the artists of the 406th Medical Laboratory, U.S. Army, in Tokyo, Japan, and by Thomas M. Evans, Linda Heath, and Niphan Ratanaworabhan of our staff. Original drawings for this bulletin were prepared by Gloria Gordon of our staff and the senior author. The wing photographs were made by Sally Craig of the Walter Reed Army Institute of Research, Washington, D.C., and Jack Scott of the Smithsonian Institution. To all these persons we extend our thanks.

This investigation was supported in part by U.S. Army Medical Department Contract No. DA-49–193– MD-2177.

III

The first species of Culicoides reported from the West Indies was described from the beaches of northern Cuba in 1851 by Felipe Poey, "the father of Cuban Zoology," as Oecacta furens (fig. 1). Poey noted the abundance and bloodsucking attacks of "el jejen" on the Cuban coasts and speculated that it must be breeding in the mangroves or marshes along the seashore.

In the English speaking islands of the Antilles these pests are known as "sandflies." Williston (1896) 1 described three species of Culicoides from Saint Vincent as Ceratopogon maculithorax, C. decor, and C. phlebotomus. H. H. Smith, who collected naterial for Williston, added the note on C. phlebotomus: "This is the common 'sand-fly' about the southern end of the island."

Coquillett (1901) described Ceraopogon melleus from Florida; this loodsucking species is also a pest in he Bahamas. Little was added to ur records of West Indian sandflies

The year in italic after the authors' ames indicates the reference in Literaire Cited, p. 86.

until Edwards (1922) (1922) described Culicoides loughnani and a variety jamaicensis from Jamaica. Hoffman (1925) reviewed the North American, Central American, and West Indian Culicoides and described C. trinidadensis.

In 1942 Fox began a concentrated study of the Caribbean Culicoides resulting in a long series of papers (Fox, 1942, 1946, 1947, 1948, 1949, 1952a, 1952b, 1955a, 1955b; Fox and Hoffman, 1944; Fox and Kohler, 1950; Fox and Maldonado, 1953; Fox and Garcia-Moll, 1961; and Kohler and Fox, 1951).

Beck (1951) began her study of Florida Culicoides with the description of a new species, C. floridensis, which also occurs in the Bahamas.

In a far-sighted and effective response to the difficult sandfly pest problem at resort hotel and beach areas on the north coast, the Jamaican Government established a Sandfly Research and Control Laboratory under the Ministry of Health at Montego Bay in 1959. Since then this laboratory has contributed immensely to our knowledge of the biology and control of West Indian

1

sandflies, beginning with the work of D. S. Kettle and later that of J. R. Linley and John B. Davies (see Literature Cited for their contributions). A similar laboratory has been established on Grand Cayman Island and staffed by M. E. C. Giglioli and John E. Davies.

The Bahamian Government was never able to establish a coordinated

laboratory such as these, but in the early 1930's it enlisted the help of J. G. Myers, a government entomologist, from Trinidad. Since 1950 it has relied heavily on the assistance of private entomologists from the United States, two of whom should be mentioned for their important work on the biology and control of Bahamas sandflies-Edwin A. Sea

brook of the Palm Beach County, Fla., Mosquito Abatement District, and Glenn M. Stokes of New Orleans, La.

Setting the geographic limits of the West Indies is slightly arbitrary and controversial depending on the purposes of the study. Geological, biological, and political considerations result in different limits. For the purpose of this study, we have excluded Trinidad and Tobago from the West Indies because these islands have a sandfly fauna essentially the same as the adjacent Venezuelan mainland and because a review of their sandfly species is being reported elsewhere. We will refer occasionally to Trinidad as "West Indian" when a few Antillean species are found there but not on the South American mainland. Although the Dutch island of Aruba falls within the same category as Trinidad in being closely related to the Venezuelan

mainland in its biogeography, we have included its only known Culicoides species in our review because it has been neglected in other reports. The Bermuda Islands share some Culicoides species with the Southeastern United States and only one with the West Indies. We are excluding them from our study and we refer the reader instead to the reports of Williams (1956, 1957) and Wirth and Williams (1957).

Regrettably our coverage of the West Indies has been extremely uneven, and today we know practically nothing of the Culicoides of the important islands of Cuba and Hispaniola and most of the islands of the Lesser Antilles except Dominica. Perhaps one of the major accomplishments of this bulletin will be to point out these gaps in our knowledge and to provide a tool for filling them in when field work can be done in these neglected areas.

Our knowledge of the biology and biting habits of Culicoides is still so fragmentary that we have scarcely any indication of their importance as vectors of pathogenic organisms, but their role is probably minor as compared with that of mosquitoes, fleas, lice, and ticks. However, Culicoides may have an important role in the transmission of some diseases, mainly as vectors of filarial worms (Sharp, 1927, 1928; Steward, 1933; Buckley, 1933, 1934, 1938; Dampf, 1936; Henrard and Peel, 1949; Romaña and Wygodzinsky, 1950; Chardrome and Peel, 1951; Hopkins,

1952; Hopkins and Nicholas, 1952; Mirsa et al., 1952; and Duke, 1954, 1956). These species may also transmit certain groups of viruses such as bluetongue of sheep and cattle (du Toit, 1944; Price and Hardy, 1954; Foster et al., 1963; Bowne et al., 1964, 1966); horsesickness (du Toit, 1944); buttonwillow virus (Reeves et al., 1970); malarialike protozoa including Haemoproteus of birds (Fallis and Wood, 1957; Fallis and Bennett, 1960, 1961); Leucocytozoon of chickens (Akiba, 1960); and Hepatocystis of monkeys (Garnham et al., 1961).