Q 3 U563 CRLSSI SMITHSONIAN CNS PEL UT tO N MUSEUM OF NATURAL HISTORY UNITED STATES NATIONAL MUSEUM BULLETIN 283 Revision of the Oriental Species of Stilobezzia Kieffer (Diptera, Ceratopogonidae) SUJIT KUMAR DAS GUPTA Hooghly Mohsin College, Chinsura, West Bengal, India WILLIS W. WIRTH U.S. Department of Agriculture, Washington, D.C. SMITHSONIAN INSTITUTION PRESS WASHINGTON, D.C. 1968 Publications of the United States National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States National Musewm and United States National Museum Bulletin. In these series are published original articles and monographs dealing with the collections and work of the Museum and setting forth newly acquired facts in the field of anthropology, biology, geology, history, and technology. Copies of each publication are distributed to libraries and scientific organizations and to specialists and others interested in the various subjects. The Proceedings, begun in 1878, are intended for the publication, in separate form, of shorter papers. These are gathered in volumes, octavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, separate publications consisting of monographs (occasionally in several parts) and volumes in which are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the botanical collections of the Museum have been published in the Bulletin series under the heading Contributions from the United States National Herbarium. This work forms number 283 of the Bulletin series. FRANK A. TAYLOR Director, United States National Museum U.S. GOVERNMENT PRINTING OFFICE WASHINGTON : 1968 For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Price 70 cents (paper cover) Introduction This paper is the second of what we hope will be a series of revisions of Oriental Ceratopogonidae, the first being a revision of the Oriental Alluaudomyia by Wirth and Delfinado (1964). The ceratopogonid material which has been sorted and mounted on slides from extensive light trap collections brought to the U.S. National Museum for the study by Wirth and Hubert (in preparation) on the Culicoides of southeast Asia provides a source which we hope will facilitate badly needed revisions of other genera of biting midges. This study is based on a total of 2,617 specimens of which 1,192 are mounted on slides. Types, including paratypes, were selected only from slide-mounted material; the ‘‘Additional Specimens” in our lists are also mounted on slides unless noted as “pinned” or “‘alcohol.’’ In “Distribution” the type locality is given first, followed by a semicolon; then the other localities are listed alphabetically or broadly by general distribution patterns in the case of wide-ranging species. In the locality data lists, the arrangement is alphabetical as far as possible, with the country first (CAPITALIZED), then the State or Province, district or town, etc. The English transliteration of Thailand localities has been done as far as possible according to the Times Atlas of the World. For their generous cooperation in collecting and sending us Oriental material, we are especially indebted to the following persons and institutions: H. C. Barnett, A. A. Hubert, H. E. McClure, and R. Traub, associated with the Walter Reed Army Institute of Research and the Medical Research Institute at Kuala Lumpur, Malaya; D. H. Colless and R. H. Wharton, Commonwealth Scientific and Industrial Research Organization, Canberra, Australia; M. E. Griffith and D. R. Johnson of the Agency for International Develop- ment, Washington, D.C.; J. L. Gressitt, T. C. Maa, L. W. Quate, and C. Yoshimoto of the B. P. Bishop Museum, Honolulu; and J. E. Scanlon and Manop Rattanarithikul formerly of the SEATO Medical Research Laboratory, Bangkok. The types of our new species are deposited in the U.S. National Museum in Washington and the B. P. Bishop Museum in Honolulu. Paratypes, when available, will be deposited in the Bishop Museum, Honolulu; British Museum (Natural History), London; Indian Museum, Calcutta; School of Public Health, University of Sydney; 1 a U.S. NATIONAL MUSEUM BULLETIN 283 and the Commonwealth Scientific and Industrial Research Organi- zation, Canberra, Australia. This study was made possible by a grant from the United States Educational Foundation to the senior author to permit study at the U.S. National Museum from January until August 1965. Classification A systematic list of the 57 known Oriental Stilobezzia species is presented in table 1. We can arrange these species conveniently into three subgenera, Stilobezzia, Hukraiohelea, and Neostilobezzia. Within these subgenera, only the first two have lent themselves to a satis- factory breakdown into species groups; most of the Neostilobezzia, at least the Oriental fauna, defy attempts at natural arrangement. The Brevicostalis Group of the subgenus Eukraiohelea is unique among the Stilobezzia species in having a shortened costa, a stout spine on the trochanter, and an extremely small, lappetlike aedeagus. In the subgenus Stilobezzia, the Boharti Group possesses several neostilobezzian features in having a spacious first radial cell, long costa, and little difference in the lengths of tarsomeres III and IV. The species of the Poikiloptera Group have some eukraiohelean features such as the extreme reduction of the first radial cell, the long, columnar parameres, and linear aedeagal sclerites with their prominent dorsomedian lobes that suggest that Hukraiohelea and Neostilobezzia are only distantly related to each other though related separately to Stilobezzia sensu stricto. In the appendix we have been only partially successful in assigning the described world species of Stilobezzia to proper subgenera and species groups. In the subgenera EHukraiohelea and Stilobezzia, our eight species groups are well contrasted on the basis of color patterns and body setation as well as by internal genitalic structures, and we are fairly confident of their natural basis. When it comes to the subgenus Neostilobezzia, however, the problem of species groups is quite different. Except for those belonging in the Speculae Group, few group-level similarities could be found in the Oriental species we studied, almost each being novel in its own way. These species have, as a rule, no significant color pattern on their body parts, and the setation, including such a basic subgeneric criterion as the surface macrotrichia of the wing, presents such devious intraspecific variation that even male-female collation from random collection is often difficult. Further study of both sexes of many of the described species and more detailed information on their distribution will be necessary in order to arrive at a natural group classification of this subgenus. 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Because the descriptions do not give the present-day diagnostic characters and the type material has not been available to us for study, we have been unable to place satisfactorily the following Stilobezzia species described from the Oriental Region: bimacula (Kieffer) from India, a member of the Notata Group; castanea Macfie from Sumatra and Malaya (Macfie described the species twice, once (1934, p. 222) from Sumatra, and again (1934, p. 284) from Malaya; it belongs to the subgenus Neostilobezzia, but the female cannot be identified specifically with any of our Malayan species.); crassipes Kieffer from India, in the Notata Group; decora Kieffer from Taiwan, in the Notata Group; minima Kieffer from India, possibly in the Brevicostalis Group; perspicua Johannsen from Sumatra, in the Notata Group; soror Johannsen from Bali, subgenus Neostilobezzia; and viridiventris (Kieffer) from Burma, unplaced here. Terminology For a detailed account of ceratopogonid anatomy and for an explanation of the terminology used in taxonomic descriptions in this family, the reader is referred to the general accounts of Carter, Ingram, and Macfie (1920), Edwards (1926), Lee (1948), Wirth (1952), and Tokunaga and Murachi (1959). We would like to explain several details of terminology and usage in this paper that might otherwise cause confusion. Antenna: In the females of Ceratopogonidae, the 3 primary seg- ments of the Dipterous antenna—scape, pedicel, and flagellum— form 15 divisions or antennomeres that we number I to XV in our descriptions. The first (1) is the ringlike scape more or less hidden in the head capsule; the second (II) is the large and globular pedicel; the distal 13 divisions (IIJ—-XV) form the flagellum of which the distal 5 usually differ in length and structure from the preceding 8. The antennal ratio (abbreviated AR in our descriptions) is the value obtained by dividing the combined lengths of the distal 5 by the combined lengths of the preceding 8 flagellomeres. Wing: Wing length is measured from the basal arculus to the wingtip; the costal ratio (CR) is the value obtained by dividing the length of the costa, measured from the basal arculus to the end of the second radial cell, by the wing length. In our descriptions, the first and second radial cells are abbreviated 1RC and 2RC, respectively. Legs: On the hind legs, the tibia bears at the distal end a blunt spur on the flexor side and an oblique row of long tibial spines. The tarsal ratio (TR) is the value obtained by dividing the length of the hind basitarsus by the length of the second hind tarsomere. Abdomen: Length of the spermatheca is obtained by measuring from the tip of the sclerotized portion of the neck to the apex of the spermatheca. In the male genitalia, we are continuing the traditional use of the term ‘“‘paramere’’ for the internal sclerotization between the aedeagal sclerotization and the base of the basistyle, though we are aware of the anatomists’ arguments that this term could be applied better to the combined basistyle-dististyle and the term “claspettes” applied to the internal structure. In the genus Stilobezzia, the term “submedian lobes” seems preferable for the structures that are ordi- narily called “apicolateral processes” in other genera and sometimes even in this genus. Likewise, the processes of the basal portion of the 9 10 U.S. NATIONAL MUSEUM BULLETIN 283 basistyle, which, in older terminologies based on Culicoides, have been called the dorsal and ventral roots of the basistyle, require special terminology as explained in the generic diagnosis. The illustrations were made by the senior author who normally used an ocular grid and squared paper; a few (i.e., thoracic patterns) were freehand sketches from pinned specimens. No attempt was made to preserve a uniform scale. A careful selection of parts was made to illus- trate a species. Within a group, the more common or better known species is selected for detailed illustration, and in other species of that group, only the diagnostic aspects are figured. Thus, the interorbital region, maxillary palpus, femora-tibiae, and last 2 tarsomeres and claws of the female are ordinarily illustrated for only 1 species of each group. Details of the male and female genitalia and the female spermathecae have special taxonomic significance; for this reason, they are figured whenever possible. Our measurements were made with an ocular micrometer on speci- mens cleared in phenol and mounted on slides in balsam. The measure- ments are usually presented in the descriptions in the form “mean (minimum-maximum, n = number of measurements)” unless a single specimen is being described. Characters with unit values such as man- dible teeth and tibial spines are given as ‘“‘range (average, n = number of specimens counted).”’ Taxonomic Treatment Genus Stilobezzia Kieffer (See detailed illustrations: figures 45, 47, 71, 80; for synonymy, see citations under subgeneric headings.) GENERIC DIAGNOSIS.—Species usually of moderate to large size, slender, with body hairs not numerous. Head: Eyes bare, their inner margins nearly contiguous to broadly separated; interorbital bridge gently curved, bearing a seta in front (female) or shaped as inverted V with no seta in front (male). Female antenna (figs. 1-40) with I[I—X oval to barely cylindrical with smooth contour, and XI—XV strongly cylindrical with rugose contour. Male antenna with well-developed plumes; XIII—XV long and strongly cylindrical with rugose contour, III—XII oval to a little elongated with vasiform base. Female mandible abruptly tapering on distal third with few strong teeth on inner side; often with fine toothlike serrations on outer margin near apex. Maxillary palpus in both sexes with III and V subequal to slightly unequal, III with short to long sensilla usually borne in a small, round, shallow, subapical sensory pit. Thorax: With or without color pattern, anterior margin of scutum evenly rounded or sometimes narrowed and more convex with a distinct median tubercle. Humeral pits present but inconspicuous; transverse suture distinct laterally. Legs: Slender, unicolorous, or ornamented with definite color markings usually indicative of group relations. Claws of female usually large, strongly unequal, sometimes appearing as a single claw with a subbasal tooth (which we interpret as a specialized feature produced by reduction of the small claw and its more com- plete fusion with the larger); male claws equal (rarely unequal), nearly always with tips bifid a very short way; no visible empodium. Tarsomere V in female unarmed (as in the male) or armed with strong spines sometimes modified into specialized, long, blunt-tipped batonnets, usually 2 in number, rarely more. Tarsomere IV bilobed or cordate distally, sometimes the lobes armed with spines; I-III variably bearing stout, usually dark, sharp spines ventrally, usually distally but sometimes at base and midportions. Tarsomeres I—II of all legs beset with ventral rows of uniformly smaller thorny setae (bulbous setae of Kieffer, Clastrier, etc.) usually in 1 row per tarsomere on fore and midlegs, in 2% to 3 rows on I, and 2 rows on II on hind legs. 11 11S U.S. NATIONAL MUSEUM BULLETIN 283 CS Eat 7 rE = See re aba oe ON non kPwWND = 11 V2 13 14