I 30 April 1981 PROC. ENTOMOL. SOC. WASH. 83(2), 1981, pp. 339-358 THE TAB ANUS STRIATUS COMPLEX (DIPTERA: TABANIDAE): A REVISION OF SOME ORIENTAL HORSE FLY VECTORS OF SURRA1 JOHN F. BURGER AND F. CHRISTIAN THOMPSON (JFB) Department of Entomology, University of New Hampshire, Dur- ham, New Hampshire 03824; (FCT) Systematic Entomology Laboratory, IIBIII, Agric. Res., Sci. and Educ. Admin., USDA, %U.S. National Mu- seum of Natural History, Washington, DC. 20560. Abstract.?Three distinct species previously confused with and called Tabanus striatus are characterized: Tabanus striatus Fabricius in the north- ern and western part of the Oriental Region; T. partitas Walker in the eastern and southern part of the Oriental Region and Micronesia; and T. triceps Thunberg on the Indian subcontinent. Illustrations and a key are given, along with a review of the previous literature on these bloodsucking pests and vectors of livestock diseases. Three species of Tabanus in the Oriental Region with trivittate abdomen (Tabanus striatus Fabricius, Tabanus partitus Walker and Tabanus triceps Thunberg) have, since their description, been subject to misinterpretation by many authors. One of the species previously called striatus has been incriminated as a vector of surra, an important disease of horses. Many of the fundamental studies on this species and others in the complex have been published under incorrect names. Determination of correct synonymy and application of correct names have not been possible with confidence due to inaccessibility of types, a myriad of synonyms incorrectly placed, and crit- ical morphological characters misconstrued or ignored. Burton (1978) aptly termed the problems associated with these species "chaotic." To resolve these taxonomic problems, their history is reviewed. Complete synonymy, diagnosis, history, and distribution are given for each species. We recognize three distinct taxa previously confused with and called Ta- banus striatus: Tabanus striatus Fabricius in the northern and western part of the Oriental Region, from Pakistan, India, and Sri Lanka to China; T. 1 Scientific Contribution Number 1066 from the New Hampshire Agricultural Experiment Station. Purchased by United States Department "of'-A-g-ri cul tur<= f or Off icial use. *? ?wsw^sw" 340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON partitus Walker from Thailand (and possibly Burma), Malaysia, Indonesia, Philippines, and Micronesia; and T. triceps Thunberg from Sri Lanka, India, and Pakistan. These species can be separated by the following key: 1. Females (eyes separated) 2 - Males (eyes contiguous) 4 2. Usually larger species (14-16 mm); foretibia uniformly orange to orange brown, not noticeably darkened apically (Fig. 3); frontal cal- losity with basal portion long and narrow, narrowly separated from eye margins ventrally and receding from eye margins dorsally (Fig. 1); abdominal venter uniformly gray tomentose and pale pilose, lack- ing a median dark stripe (Fig. 7) triceps Thunberg - Usually smaller species (10-13 mm); foretibia sharply bicolored, pale on basal %, blackish on apical Vs (Fig. 6); frontal callosity with basal portion contiguous with eye margins for most or all its length (Fig. 2); abdominal venter with distinct broad median dark stripe (Fig. 8) 3 3. Abdomen with dorsal median pale stripe evanescent or absent on tergum 2 (Fig. 5); abdominal ground color blackish striatus Fabricius - Abdomen with dorsal median pale stripe complete, fully developed on tergum 2 (Fig. 4); abdominal ground color dark brown to brown black partitus Walker 4. Foretibia uniformly orange to orange brown (Fig. 3); abdominal ven- ter uniformly pale yellowish white to yellow tomentose and pilose (Fig. 7) triceps Thunberg - Foretibia bicolored, pale basally, becoming blackish on apical 1/4 (Fig. 6); abdominal venter yellowish white tomentose with a broad dark midstripe (Fig. 8) 5 5. Costal cell clear, never tinted striatus Fabricius - Costal cell yellow tinted . partitus Walker Tabanus striatus Fabricius Tabanus striatus Fabricius, 1787: 356. Type-locality: China. Lectotype UZMC. Subsequent references: Surcouf, 1923: 196 (taxonomy); Isaac, 1924b: 108 (biology, immature stages); Chvala and Lyneborg, 1970b: 546 (lectotype designation); Stone, 1972: 639 (taxonomy), 1975: 70 (catalog citation); Burton, 1978: 71 (taxonomy, Laos, Thailand distribution rec- ords, biology). Tabanus hilaris Walker, 1850: 49. Type-locality: East India. Holotype male BM(NH). Subsequent references: Bigot, 1891: 269, van der Wulp, 1896: 60, Kertesz, 1900: 53, 1908: 249 (catalog citations); Ricardo, 1911: 153 AV ?s*?sww%T?v??r VOLUME 83, NUMBER 2 341 (taxonomy), 1916: 407 (Hong Kong); Fletcher, 1916 (life history), 1917 (oviposition). Tabanus tenens subform cambodiensis Toumanoff, 1953: 201. Type-local- ity: not stated (Cambodia). Holotype lost. Tabanus striatus (in part); Bigot, 1891: 268, van der Wulp, 1896: 58 (catalog citations); Kertesz, 1900: 71, 1908: 281 (world catalogs); Ricardo, 1911: 149 (taxonomy); Fletcher, 1916 (life history, surra vector), 1917 (ovipo- sition); Austen, 1922a: 445 (taxonomy); Schuurmans Stekhoven, 1926: 63, 1928: 438, 1932a: 65 (taxonomy, distribution); Senior-White, 1927: 51 (catalog citation); Wu, 1940: 186 (catalog citation); Philip, 1959: 606, 1960: 57, 1973: 60 (taxonomy, synonymy, distribution); Chvala and Lyneborg, 1970a: 365 (taxonomy, distribution); Moucha, 1976: 152 (world catalog). erroneous citations to striatus (in total or part): van der Wulp, 1880: 163, 1881: 16, 1885: 71, 1896: 58 (includes also dorsilinea and partitus); Bigot, 1891: 208 (includes also dorsilinea and partitus); Kertesz, 1900: 71, 1908: 281 (includes also dorsilinea and partitus); Ricardo, 1911: 149 (includes also triceps and partitus); Mitzmain, 1913 (refers to partitus); Fletcher, 1916, 1917 (may also include triceps); Krober, 1924: 18 (refers to parti- tus); Schuurmans Stekhoven, 1926: 63, 1928: 438, 1932a: 65 (includes partitus); Senior-White, 1927: 51 (includes partitus and triceps); Nie- schulz, 1926a-c, 1927a-c, 1928, 1929a-b, 1935a-b, 1936, Nieschulz and Ponto, 1927 (refers to partitus); Kelser, 1927 (refers to partitus); Wu, 1940: 186 (includes triceps and partitus); Philip, 1959: 606, 1960: 57, 1973: 60 (includes partitus); Stone, 1960: 52 (refers to partitus); Chvala and Lyneborg, 1970a: 365 (includes partitus); Moucha, 1976: 142 (includes triceps and partitus). Diagnosis.?Tabanus striatus Fabricius is closely related to partitus. Bur- ton (1978) provided characters to separate the two in Thailand (as striatus and megalops), and these can be applied elsewhere in the range of both species. In particular, females of striatus do not have a pale tomentose and pale haired midstripe on the second tergum. These characters will separate it from both partitus and triceps females that do not have an abbreviated midstripe. The ground color of the dorsal abdominal surface is black in striatus females, dark brown in partitus. Males of both striatus and at least some partitus have the midstripe abbreviated, i.e. absent or evanescent on the second tergum, but the costal cell is completely hyaline in striatus and yellow tinted in partitus. Other differences were noted between striatus and partitus, but these may not hold up when more material is examined from intervening localities. In females, the apical segment of the palpus is slightly shorter, more stout and less yellowish in striatus than in Philippine partitus; the scutellum has the black pilosity (pale in partitus); the prescutellar scler- ite has black pilosity (pale in partitus); the foretibia is less extensively or- ev&&^j&*&?*--&&^'^ 342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. Head, frontal view. 3, 6. Foreleg, lateral view. 4-5. Abdomen, dorsal view. 7- 8. Abdomen, ventral view. 1,3,7, Tabanus triceps. 2, 5, 6, 8, T. sliiulus. 4, T. partitus. VOLUME 83, NUMBER 2 343 ange basally, with apical fourth black; the stem of the halter is pale brown (yellowish white in partitus); sublateral pale stripes on abdomen are not conspicuously jagged or steplike (distinctly jagged or steplike in partitus (see Figs. 4, 5)); the venter is gray tomentose (more yellowish gray in par- titus). The male of striatus always lacks the midstripe on the second ab- dominal tergum, although traces of pale pilosity may be present. The male of partitus has the midstripe on the second abdominal tergum variably de- veloped, being nearly absent on some specimens, or being present and crossing the tergum to greatly narrowed in others. History.?Tabanus striatus was originally described from China by Fa- bricius (1787). A specimen in the Zoological Museum, Copenhagen was designated as lectotype by Chvala and Lyneborg (1970b). Since its descrip- tion, this species has been frequently interpreted too broadly, usually in- cluding one or more additional species, most commonly a southern "variant form" from Thailand, Malaysia, Indonesia, and the Philippines, now known as Tabanus partitus Walker. Confusion about the limits of striatus has been discussed by Burton (1978). Tabanus striatus and partitus (Java distribution records) were combined in the work of van der Wulp (1880, 1881, 1885) and the catalogs of Bigot (1891), van der Wulp (1896), and Kertesz (1900, 1908), but the greatest source of confusion about the limits of striatus was in Ricardo's (1911) work. She included both triceps of India (then called tenens Walker) and partitus under striatus. Her concept of striatus was hopelessly confused, as later stated by Austen (1922a). Ricardo incorrectly gave Fabricius (1794) as the original citation and "China and Java" as the type-localities of stria- tus. Ricardo even separated Tabanus hilaris Walker, a synonym of striatus, from striatus by the presence of an abbreviated median abdominal stripe, the character Fabricius used to define striatus and which Ricardo quoted in her paper. Most earlier authors (van der Wulp, Bigot, Kertesz, Ricardo, and Wu) also included the name dorsilinea Wiedemann as a synonym of striatus, but, as shown by Burton (1978: 78), this name is the senior synonym for a species previously called bicallosus Bigot (Ricardo, 1911: 129) or macer Bigot (Austen, 1922b: 264; Senior White, 1927: 44). Austen (1922a) distinguished triceps (as tenens) from striatus, but com- bined partitus with striatus, an interpretation followed by most subsequent workers, including Schuurmans Stekhoven (1926, 1928, 1932), Senior-White (1927), Nieschulz (all papers), Wu (1940), Oldroyd (1957), Philip (1959, 1960, 1973), Stone (1960) and Moucha (1976). Their use of characters, particularly of the legs and abdomen, did not allow differentiation of these two species {partitus and striatus). Austen (1922a) described striatus as having the midstripe on the second abdominal tergum more or less obsolete, at least much less distinct than on 344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON the following terga. He also stated, however, that specimens of striatus, especially those which have the midstripe not obliterated on the second tergum, are liable to be mistaken for examples of triceps (as tenens). Austen thus included specimens that do have a midstripe on tergum two (i.e. par- titus) as well as those that do not in his concept of striatus. Krober (1924), discussing "striatus" from the Philippines, also mentioned that the mid- stripe on the abdomen was not always clear, probably a reference to male partitus, a species that sometimes has the midstripe evanescent on the sec- ond segment. Surcouf (1923) was the first author to distinguish three distinct taxa throughout the geographic range of the striatus complex. He correctly rec- ognized striatus Fabricius as distinct from another species in India that he named strophiatus, since he mistakenly believed tenens Walker to be a synonym of striatus. He did, however, correctly associate hilaris Walker with striatus. Surcouf clearly separated striatus from related taxa by re- stricting striatus to those specimens with the median abdominal stripe be- ginning on the anterior margin of the third tergum. For specimens with the midstripe beginning on the anterior border of the second tergum, Surcouf recognized two species, one he correctly called partitus Walker with dark legs, with ruficallosus (lapsus for rufocallosus Bigot) and manilensis Schi- ner as synonyms, and strophiatus Surcouf with testaceous legs, a new name for striatus of authors from India, China, and Indochina. Surcouf s inter- pretation of these taxa and their names was accurate except for his mistaken synonymy of tenens Walker under striatus and the renaming of the true tenens (=triceps) as strophiatus. He did correctly recognize three distinct taxa, utilizing characters that will separate these taxa. Unfortunately these characters were not consistently applied by subsequent authors, who over- looked Surcouf s work. Schuurmans Stekhoven (1926) recognized a northern, "typical" form of striatus, and a southern form that showed deviations from a specimen from Punjab, India that was compared with the type of striatus by K. L. Hen- driksen. Schuurmans Stekhoven found that females caught in "southern areas" had wings with a yellow-brown costal cell and the midstripe paler but not abruptly interrupted on the second tergum. The males had wings with a yellow-brown costal cell. Thus, Schuurmans Stekhoven recognized differences that today are used to separate striatus from the related partitus, but he did not recognize these "forms" as distinct species. The use of the name striatus in all of Nieschulz's fundamental work on surra and its transmission refers to partitus as his studies were restricted to Indonesia, principally Java. Philip (1959) examined the types of hilaris Walker, partitus Walker and rufocallosus Bigot and concluded that all were variants of striatus. He also found that the type of manilensis Schiner did not disagree significantly from VOLUME 83, NUMBER 2 345 the type of striatus except for the distally brown foretibia. Philip discussed the variable nature of the middle abdominal stripe on tergum two. He noted that the midstripe was either interrupted on tergum two in many Philippine males (but in only one female) or narrowly extended across the second tergum in "many other males" and most females, with one female showing the stripe "hardly narrowed." Although this character would more closely apply to triceps than striatus, Philip considered it to be variable throughout the broad range of striatus in the Oriental Region. Isaac (1924b) discussed the life history of striatus in India. His illustra- tions of the male and female show clearly that he was working with true striatus. Stone (1972) studied the types of several supposed synonyms of striatus and found that two distinct species were involved. He believed that the original description of striatus more closely agreed in abdominal coloration with specimens from China, its type-locality, than with specimens called striatus by earlier authors from more southern portions of the Oriental Re- gion. He separated triceps (as tenens) from striatus by the strong middorsal pale stripe present from tergum I-VII, palpus longer and less swollen ba- sally, and with the abdomen yellow brown rather than black brown. Stone believed the complete slender midstripe of partitus ruled out conspecificity with striatus and triceps (as tenens). He thereby implied that partitus was a distinct species. Although this character is not reliable for both sexes of partitus, Stone was the first author since Surcouf to recognize partitus as distinct from striatus. However, he placed manilensis and rufocallosus as synonyms of triceps (as tenens), rather than partitus, and did not give any taxonomic characters to support his synonymy. Moucha (1976) did not follow Stone's interpretation, but reverted to Phil- ip's concept of striatus, including under it most of the synonyms of partitus, as well as strophiatus Surcouf. He did, however, correctly synonymize tenens subform cambodiensis Toumanoff under striatus. Burton (1978) has completely described striatus and thoroughly discussed the confusion of striatus with closely related partitus (as megalops), men- tioning that the abdominal striping pattern will not serve to distinguish males of striatus and partitus. Herein lies at least some of the confusion previous authors encountered in attempting to define the limits of striatus, as the midstripe can be interrupted in males of partitus as well as striatus. How- ever, as Burton pointed out, the yellow tinted costal cell in the male of partitus will separate it from striatus males which have a hyaline costal cell. Interestingly, this character was mentioned by Schuurmans Stekhoven (1926) in discussing the "southern form" of striatus, but was not considered by subsequent authors. Distribution.?Pakistan, India, Sri Lanka, northern and eastern Thailand, Laos, Cambodia, Vietnam, China. 346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Specimens examined.?30. INDIA: Madras; Nedungadu; Tanjore Dist.; Sohawa; Jhelum; Karum Bagarum, Assam: Chabua; Dinjan; Doom Dooma. SRI LANKA: 10 localities throughout the island. PAKISTAN: Lahore. THAILAND: Loei. VIETNAM: "Tonkin." Tabanus partitas Walker Tabanus partitas Walker, 1856: 9. Type-locality: Singapore. Holotype fe- male BM(NH). Subsequent references: Bigot, 1891: 270, van der Wulp, 1896: 60, Kertesz, 1900: 64, 1908: 268 (catalog citations). Tabanus manilensis Schiner, 1868: 84. Type-locality: Philippines, Manila. Holotype female NMW. Subsequent references: van der Wulp, 1896: 61 (as manillensis), Kertesz, 1900: 64, 1908: 259 (catalog citations). Tabanus rufocallosus Bigot, 1892: 197. Type-locality: Java. Holotype fe- male BM(NH). Subsequent references: van der Wulp, 1896: 63, Kertesz, 1900: 69, 1908: 276 (catalog citations). Tabanus tenens (in part): Austen, 1922a: 445 (taxonomy); Senior-White, 1927: 53 (catalog citation); Schuurmans Steknoven, 1928: 438 (Philip- pines); Wu, 1940: 187 (catalog citation); Oldroyd, 1957: 59 (taxonomy); Stone, 1975: 71 (catalog citation). Tabanus striatus (in total): Mitzmain, 1913 (biology; surra transmission in the Philippines); Krober, 1924 (Taxonomy); Nieschulz, 1926a, 1926b (bi- ology), 1926c (breeding sites), 1927a, 1927b (hymenopterous parasites), 1927c, 1928 (anthrax transmission), 1929a, 1929b, 1935a (distribution and abundance), 1935b (larval development), 1936 (biology, description of eggs, larva, and pupa); Nieschulz and Ponto, 1927 (most papers refer to surra transmission studies unless otherwise noted); Nieschulz and Kra- neveld, 1929 (haemorrhagic septicaemia of water buffalos, transmission); Kelser, 1927 (surra transmission); Schuurmans Stekhoven, 1932b: 14 (Su- matra); Stone, 1960: 52 (taxonomy, distribution in Micronesia). Tabanus striatus (in part): Ricardo, 1911: 149 (taxonomy); Krober, 1924: 18 (taxonomy); Schuurmans Stekhoven, 1926: 63, 1928: 438, 1932a: 65 (tax- onomy, distribution); Senior-White, 1927: 51 (catalog citation); Wu, 1940 (catalog citation); Philip, 1959: 606, 1960: 57, 1973: 60 (taxonomy); Chvala and Lyneborg, 1970a: 365 (taxonomy); Moucha, 1976: 142 (world catalog). erroneous citations as megalops (in part): Stone, 1972: 639 (taxonomy); Burton, 1978: 74 (taxonomy, synonym, Thailand records, biology, in- cludes triceps). Diagnosis.?Tabanus partitus was redescribed in comparison with stria- tus by Burton (1978). Female partitus are separated from striatus by the presence of a midstripe on the second abdominal tergum (absent in striatus), and male partitus have a yellow tinted costal cell (hyaline in striatus). Also, the general abdominal coloration of partitus is paler, brown to gray, than VOLUME 83, NUMBER 2 347 in striatus (black). These characters hold for both species throughout their ranges. Comparison of Burton's specimens of partitus (Cornell University Collection, courtesy of L. L. Pechuman) with those from the Philippines revealed differences as follows: The frons is narrower in Philippine speci- mens, index 1: 6.0-7.5 (1: 4.3-5.6 for Thai specimens); the apical segment of the palpus is slightly more yellowish and stouter than in Thailand spec- imens; the mesonotum is darker gray; the disc of the scutellum lacks yel- lowish tones seen on Thai specimens; mid- and hindfemora are blackish distally; the stem of the halter is brown black (yellow white in Thai speci- mens). Males of partitus from the Philippines have no yellowish tint on the scutellum as do those from Thailand. History.?Tabanus partitus was described by Walker (1856) from a female from Singapore. Since its description, it has most commonly been consid- ered a variant form of Tabanus striatus Fabricius, with striatus of authors considered to be a rather variable and widely distributed Oriental species. Various authors differentiated striatus from triceps (as tenens) found in India and Sri Lanka. Other authors clearly separated striatus from triceps (as tenens), but considered partitus to be the same as triceps. This differ- ence in concepts was based primarily on whether characters of the head and legs were used {partitus + striatus and triceps) or whether differences in abdominal striping were considered important {striatus and triceps + partitus). Ricardo (1911) included partitus as a synonym of striatus even though she quoted van der Wulp's description verbatim (1881: 16) in which the wing of a male from Soerian, Sumatra was described as having a yellow tinted costal cell, a characteristic of partitus, not striatus. Mitzmain (1913), working on transmission of surra {Trypanosoma evansi) by horse flies in the Philippines, discussed the biology of Tabanus partitus (as striatus), including a brief description of the male and female. The wing of the female was described as transparent except for the pale brown costal and subcostal cells, a characteristic of partitus. We have examined voucher specimens of striatus sensu Mitzmain and find them to be partitus. Austen (1922a) attempted to differentiate striatus from triceps (as tenens) by the midstripe being more or less obsolete on the second abdominal ter- gum, and synonymized partitus (as megalops) under triceps (as tenens). However, the tarsal characters he used will not separate striatus from tri- ceps. Also, because he incorrectly synonymized hilaris {?striatus) with triceps (as tenens), his ability to separate these taxa is questionable. Krober (1924), in his study of the tabanid fauna of the Philippines dis- cussed partitus under the name striatus, stating that the midstripe on the abdomen was not always clear on the second tergum and citing a distribution for striatus from India to the Moluccas. Schuurmans Stekhoven (1926) clearly distinguished a northern typical ' > #i*!WSR>-i?S.,*WWff" 348 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON form and a southern form of striatus, stating that those from southern lo- calities had wings with yellow-brown costal cells and mid stripe on the ab- domen paler but not abruptly interrupted on the second segment. Both males and females of striatus have a clear costal cell, so Schuurmans Stekhoven's southern form clearly refers to partitus. In subsequent papers, Schuurmans Stekhoven (1928, 1932a, b) provided additional collection records from In- donesia, also listing the "typical" form (i.e. striatus Fabricius) from Hanoi (Vietnam) and Fukien Province (China). Nieschulz (1935b, 1936) studied the hatching and development of the im- mature stages of partitus (as striatus) in Indonesia. He also provided excellent illustrations of the larva and pupa of partitus as well as notes on the larval habitat. The illustrations are among the best available for any Oriental horse fly by early workers and allow comparison with known larvae and pupae of other species. Unfortunately, the larvae of triceps and striatus are less well illustrated and cannot be compared easily with partitus. In all of the fundamental work of Nieschulz on surra, the name striatus refers to partitus. Oldroyd (1957) accepted Austen's synonymy of partitus (as megalops) and triceps (as tenens), with triceps distinguished from striatus by the presence of three abdominal stripes, being longer and more gently tapered and the midstripe complete from front to rear. Philip (1959, 1960, 1973), however, continued to consider specimens of partitus as variants of stria- tus. His synonymy of striatus is confused because he incorrectly placed partitus Walker and rufocallosus Bigot as synonyms of striatus and partitus as a synonym of triceps Thunberg. He also stated that the type female of manilensis Schiner (=partitus) "did not differ significantly" from striatus, even though in his discussion of striatus from the Philippines he mentioned the middorsal pale abdominal stripe on the second tergum as being present in Philippine material. Stone (1960) recorded partitus (as striatus) from Guam. We have exam- ined Stone's specimens from Guam in the USNM and did not find any major differences from partitus. Stone recognized that the synonymy of striatus was confused, involving more than one species, and he clearly distinguished triceps of India from his Guam specimens by the larger, browner body, entirely brown forefemora and the venter of the abdomen lacking the dark- ened midstripe found in striatus. Stone did, however, list triceps from the Philippines, so he may have included partitus, at least in part, in his concept of triceps. Stone (1972), after examining the types of several supposed synonyms of striatus, concluded that two species were involved and that most of the synonyms did not apply to striatus. He recognized discrepancies between true striatus and the variant form from the southern Oriental Region. He VOLUME 83, NUMBER 2 349 also recognized that megalops has a complete but slender midstripe that he believed ruled out conspecificity with either striatus or triceps. Although Stone was correct in this assertion, the length of the middorsal stripe will not suffice to separate megalops from striatus complex. Further, Stone erroneously grouped synonyms of partitus with triceps (as tenens) (i.e. manilensis and rufocallosus), but he was the first author since Surcouf to recognize two distinct taxa. Also he gave triceps a much wider distribution than it really has. In the Oriental Diptera Catalog, however, Stone (1975) synonymized partitus under triceps (as tenens). Moucha (1976), unlike Stone, followed Philip's broad interpretation of striatus and placed partitus, manilensis, and rufocallosus in synonymy un- der striatus, as well as dorsilinea Wiedemann (as macer Bigot), a distinct species. Burton (1978) summarized the past confusion surrounding the synonyms of striatus and partitus (as megalops) and gave excellent taxonomic char- acters for separating the two taxa. Unfortunately, he mistakenly believed Tabanus megalops Walker from Java to be conspecific with specimens he collected in Thailand. Undoubtedly, Burton was mislead by Stone's syn- onymy. Burger has thoroughly examined the holotype male of megalops in the British Museum (Natural History) and compared it to males from Thai- land collected and determined as that species by Burton. The holotype of megalops, in fair condition, differs from the Thailand specimens in having the body stouter, costal cell clear, middorsal abdominal pattern being a series of very narrow triangles, not a parallel-sided stripe, forecoxae and femora orange brown, foretibiae uniformly brown, not bi- colored, and midventral abdominal dark stripe evanescent. Based on these differences, we believe megalops is not conspecific with Burton's Thailand specimens or with other material examined from throughout the southern Oriental Region. The earliest available name for the taxon conspecific with specimens collected by Burton and others examined by us is Tabanus par- titus Walker. The holotype of partitus is in good condition and has distinctly bicolored foretibiae, basal callosity contiguous with eyes below, pale ab- dominal middorsal stripe complete (although narrowed anteriorly) on tergum two, and distinct midventral dark stripe, all the characteristics of the south- ern component of the striatus complex. The holotype male of megalops closely resembles several males presently in the BM(NH) collection determined as Tabanus rubidus Wiedemann by Austen, Oldroyd, and others. The configuration of the abdominal triangles also is reminiscent of some rubidus males, but the triangles of the type are narrower than in the presumed rubidus males examined. The condition of the type is such that assignment to the rubidus group as defined by Burton (1978) is certain, but species assignment is difficult, especially since the 350 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON characters separating males of these species are poorly defined at present. Distribution.?Burma, Thailand, Malaysia, Indonesia, Philippines, Micro- nesia (Guam). Specimens examined.?Approximately 300. BURMA: Rangoon. THAI- LAND: Prae Nakhom Prov., Chon Bun Prov. MALAYSIA: Kuala Lumpur, Selangor. SINGAPORE. INDONESIA: Sumatra, Medan; Java, Passo- eroean, Buitenzorg. PHILIPPINES: Luzon, Leyte, Palawan, Mindanao, Osmona, Samar, Alaband, Rizel. MICRONESIA: Guam, Togcha Point. Tabanus triceps Thunberg Tabanus triceps Thunberg, 1827: 59. Type-locality: Indian subcontinent (as "Cayenne et Brasilia"). Lectotype, Zool. Inst., Univ. Uppsala. Subse- quent references: Kertesz, 1900: 74, 1908: 286 (world catalogs); Philip, 1959: 609, 1967: 1236 (taxonomy, lectotype designation), 1960: 59 (syn- onymy, Thailand record), 1970: 450 (differences from striatus Fabricius), 1973: 60 (Ceylon records). Tabanus tenens Walker, 1850: 49. Type-locality: East India. Holotype fe- male BM(NH). Preoccupied by Tabanus tenens Walker 1850, a Neotrop- ical species. Tabanus strophiatus Surcouf, 1923: 197. Type-localities: "Archipel. Indien, Chine, Indo-Chine." Syntypes, at least 56 females, MNHN, Paris. Tabanus tenens (in total): Bigot 1891: 269, van der Wulp, 1896: 60 (catalog citations); Kertesz, 1900: 73, 1908: 285 (world catalog); Austen, 1922a: 445 (taxonomy); Isaac, 1924a (male, female genitalia), 1925 (immature stages); Schuurmans Stekhoven, 1926: 163 (taxonomy). Tabanus tenens (in part): Schuurmans Stekhoven, 1928: 438 (Ceylon); Se- nior-White, 1927: 53 (catalog citation); Wu, 1940: 187 (catalog citation); Oldroyd, 1957: 59 (taxonomy); Stone, 1960: 52, 1972: 640 (taxonomy, synonymy), 1975: 71 (catalog citation). Tabanus triceps (in part): Philip, 1959: 610, 1960: 59; Moucha, 1976: 147 (world catalog). Tabanus partitus (in part): Burton, 1978: 74 (taxonomy, synonymy). Tabanus striatus (in part): Ricardo, 1911: 149 (taxonomy) erroneous citations to triceps (in part): Philip, 1959: 610 (includes partitus), 1960: 59 (includes partitus and striatus); Moucha, 1976: 147 (includes partitus). erroneous citations to tenens (in part): Senior-White, 1927: 53 (may include striatus); Schuurmans Stekhoven, 1928: 438 (includes partitus); Wu, . 1940: 187 (may include partitus and striatus). Diagnosis.?Tabanus triceps from India and Sri Lanka is quite distinct from both striatus and partitus. Thunberg's alpha syntype (lectotype) of triceps has a uniformly orange forefemur and tibia, the callosity is not con- VOLUME 83, NUMBER 2 351 tiguous with the eyes, receding from eye margins above, and the middorsal stripe on the second tergum broadly crosses that segment. It also has distinct thoracic stripes as mentioned by Philip (1959). The lectotype agrees with all the specimens we have seen from India and Sri Lanka previously deter- mined as tenens or triceps. Tabanus triceps is easily distinguished from striatus by the middorsal stripe of the abdomen distinctly crossing the second tergum in both sexes (occasionally obliterated in greased specimens), the unicolorous foretibia, the callosity narrowed above, not broadly contiguous with the eye margins, and the absence of a dark haired midventral stripe on the abdomen. The males of triceps have a yellow tinted costal cell (hyaline in striatus). Tabanus triceps differs from partitus in having the callosity narrowly separated from eye margins below and receding from eye margins above, broadly joined to the broad, squared median extension; the apical segment of the palpus slightly longer and more slender basally; the forefemur and tibia entirely orange to orange brown; the sublateral pale abdominal stripes noticeably steplike and the venter uniformly yellowish gray, lacking a broad midventral dark stripe. Males of triceps show the same differences from partitus as the females except for sex associated characters of the frons and palpus. The ground color of the abdomen of male triceps is blackish, and the middorsal stripe on the second tergum is usually well developed, while partitus males have a more reddish-brown abdominal ground color and the middorsal stripe on the second tergum is variably developed. Males of both species have a yellow tinted costal cell. Most specimens of partitus from Thailand, the Philippines, and Java are smaller than triceps from Sri Lanka and India. The mean length of partitus was 12.7 mm (range 11.2-14.3 mm) for females and 12.2 mm (range 9.6- 14.0 mm) for males. Tabanus triceps females averaged 15.0 mm (13.6-16.0 mm), while males averaged 14.4 mm (13.6-15.2 mm). The pupal aster of triceps is different from that of partitus (see figures of Isaac and Nieschulz), the latter having the lateral arms directed postero- d or sally in proximity to the dorsal arms while triceps has the lateral arms directed posteroventrally in proximity to the ventral arms. History.?Tabanus triceps Thunberg has been reported under the name Tabanus tenens Walker in most of the literature and more recently as me- galops by Burton (1978). This species generally has been considered distinct from striatus, except by Ricardo (1911). Bigot (1891), van der Wulp (1896), Kertesz (1900, 1908), Senior-White (1927), and Wu (1940) list it as a distinct species. Austen (1922a) discussed Ricardo's confusion of the two species, but did not cite detailed characters to separate triceps (as tenens) from striatus. He did mention that the description of striatus by Wiedemann 352 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON (abdomen with three unabbreviated pale stripes; femora reddish rusty brown) applied better to triceps. Surcouf (1923) used the name strophiatus for a species with the middorsal abdominal stripe beginning on the anterior border of the second tergum and the legs testaceous, a species he believed had been confused with striatus. This species was separated from a related one with dark legs that Surcouf called partitus. Thus Surcouf recognized that two species previously con- fused with striatus were distinct, one with dark legs and another with tes- taceous legs. The species he named strophiatus is the same as triceps and is synonymized with it. Isaac (1924a, 1925), in a series of papers on Indian Tabanidae, discussed the immature stages and the male and female genitalia of triceps (as tenens). Isaac's figures of the adult male and female clearly show that he was dealing with triceps. Comparison of his pupal figure for triceps with that given by Nieschulz (1935) for partitus shows that the pupal aster is distinctly differ- ent. Other features of the immature stages could not be compared due to lack of comparable figures. Schuurmans Stekhoven (1926) clearly differentiated triceps (as tenens) from striatus, but considered partitus to be a synonym of triceps (as te- nens), based on the very distinct middorsal stripe on the second segment. His description of tenens was drawn from Indian specimens and therefore refers to triceps. Philip (1959), in his study of Tabanidae from the Philippine Zoological Expedition of the Field Museum (Chicago), discussed striatus and what he considered to be its synonyms. At the same time, he discussed the status of tenens vis a vis striatus. He found that one syntype (alpha) of Tabanus triceps, described by Thunberg (1827) from "Cayenne et Brasilia" was not a Neotropical species, but was closest to tenens Walker (the Oriental species). The unpatterned eye, striped thorax, and unicolorous legs did not agree with known Neotropical species. Philip designated this alpha syntype as the lectotype of triceps, with tenens thus becoming a synonym. Philip also gave characters to separate triceps from striatus, particularly the uni- colorous legs, the uninterrupted broad mid stripe on abdominal tergum two, and the basal callosity narrowed above, well separated from the eyes. These characters would separate triceps from Philip's concept of striatus (=stria- tus + partitus), but will not adequately separate triceps from partitus. Philip (1967), in his discussion of Thunberg's species of Tabanidae from the New World, designated another lectotype, the gamma syntype, from the type-series in competition with his 1959 designation. Despite this lapse, the earlier designation has precedence, and triceps remains the valid name for Walker's tenens of the Oriental Region. Philip also stated that the alpha syntype might not have been an original syntype. However, as Thunberg described the thorax as having five white lines and the legs as completely =g VOLUME 83, NUMBER 2 353 testaceous, and because these are characters of the alpha syntype, there is no doubt that the alpha syntype was one of the specimens before Thunberg when triceps was described. Philip (1970) stated that Thunberg's gamma syntype of triceps might have come from China. However, there seems to be no reason for assuming this syntype is Oriental, based on the taxonomic characters given by Philip him- self in 1967. The gamma syntype is the same as Tabanus dorsiger var. dorsovittatus Macquart, a Neotropical species with a green and purple striped eye pattern and strongly bicolored foretibia. Females of Oriental species related to striatus have unicolorous foretibia and unpatterned eyes. The beta syntype agrees with the description and figure of Tabanus colum- bus Fairchild, another Neotropical species. This specimen also had a pat- terned eye, although the precise pattern could not be discerned due to mold- ing of the eye surface. Stone (1960) separated triceps from striatus by its larger size, the entirely brown forefemur and the absence of a darkened ventral abdominal mid stripe usually found in striatus. Subsequently, Stone (1972) reverted to tenens Walker when Philip designated a competing lectotype for triceps; thus tri- ceps became a junior synonym of dorsiger, a Neotropical species. His treat- ment of these species in the Catalog of Oriental Diptera (Stone, 1975) fol- lowed his 1972 work. Moucha, in his catalog of World Tabanidae (1976), recognized Philip's 1959 designation and used triceps as the correct name. Both Stone and Moucha, however, continued to synonymize partitus and some of its synonyms with tenens (Stone) or with both triceps and striatus (Moucha). Burton (1978) reviewed the status of partitus (as megalops), including its synonyms, in his study of the Tabanini of Thailand north of the Isthmus of Kra. He concluded that the types of tenens from East India and megalops from Java in the British Museum were conspecific, without mentioning the basis for his decision. As the type of tenens is a female and the type of megalops is a male, the association of the two may have been complicated because this would exclude use of frons characters. Burton compared the types with associated male and female specimens from Thailand. We have examined specimens from throughout the Oriental Region (including the types of megalops, partitus, rufocallosus, and tenens and a series determined as partitus by Burton) and determined that specimens from India and Sri Lanka (conspecific with the type of tenens) are specifically distinct from those collected in Thailand, Indonesia, and the Philippines (conspecific with the type of partitus). We have also had the opportunity to examine Thunberg's syntype series of triceps. The alpha syntype (lectotype) agrees with specimens from India and Sri Lanka, which were compared with the type of tenens. The uniformly yellowish-orange femora and foretibia, basal portion of the callosity reced- fKAwcwoa?!*: 354 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON ing from eyes above, and lack of darker stripe ventrally on the abdomen are particularly distinctive of the lectotype and these other specimens. This confirms Philip's (1959) determination of triceps as conspecific with tenens. Tabanus triceps thus is the correct name for the species found in India and Sri Lanka, while partitas is the correct name for the species found in Thai- land, Indonesia, the Philippines, and Micronesia. Burton also found that the name tenens for the Indian species is a junior homonym of Tabanus tenens Walker described from Brazil, also in 1850, but four months earlier. However, since triceps antedates either of Walker's names by 23 years, this homonomy does not affect the correct name of the Indian species. Distribution.?Pakistan, India, Sri Lanka. Specimens examined.?78. INDIA: Coimbatore; Karikal; Bombay; Mad- ras; Karum Bagaram; Trichinopoly; Singara, Nilgiri Hills; Tanjire Dist.; Calcutta; Kanchrapara (2 2 agree with the type of tenens Walker); Bengal; Shimuga; Mysore; Walayar Forest, South Malabar; Chinchona; Anamalai Hills. SRI LANKA: 26 localities throughout the island. UNPLACED SYNONYMS Tabanus chinensis Thunberg, 1827.?This species was described briefly by Thunberg with the habitat given both as China and the Cape of Good Hope. It possibly could be a synonym of striatus, but this is uncertain from the original description, so it is left unplaced for the present. Tabanus costalis Lichtenstein, 1796.?Bezzi (1908) first called attention to the names of Anton August Heinrich Lichtenstein. Austen (1908) dis- cussed the Tabanidae listed in Lichtenstein's Catalogus. Since only two copies of Lichtenstein's work were known to Austen, he quoted the de- scription of costalis verbatim: "295. Tabanus striatus; n. 39 (the number under which Fabricius described striatus). Item: Tabanus costalis; nobis. Taban. oculis aeneis; ferrugineus, alls hyalinis costa flava. Habitat in Co- romandel." Ricardo (1911) stated that the species is absolutely indetermin- able and should be deleted from the list of Tabanus species. Surcouf (1923) considered it a questionable synonym of striatus. Stone (1975) did not list the name in the Catalog of Oriental Diptera. Burton (1978) referred to the name under the Tabanus striatus complex, indicating its status was yet to be defined. The name appears in a sale catalog, and, thereby, may not be considered available. The arrangement of the catalog follows Fabricius (1794), and cos- talis is listed under striatus, number 295 being the number of the item in the sale catalog. The description is short, but the mention of the yellow costal cell could refer to either partitus or triceps as the males of both species have a yellow costal cell. The notation "habitat in Coromandel" K ^XX.3 BRV.JW,?p j^Ki"^T."vraj VOLUME 83, NUMBER 2 355 may refer to the Coromandel Coast of India (16?-12?N Lat.), including Mad- ras, or to Coromandel in the state of Minas Gerais, Brazil (unlikely possi- bility, Fairchild, in lift.). If the former locality is intended, then costalis may refer to triceps, but if the latter locality is intended, costalis may refer to dorsiger Wiedemann or a related species with a tinted costal cell. ACKNOWLEDGMENTS We thank Graham White and John Chainey, British Museum (Natural History), London (BM(NH)); Leif Lyneborg, Universitetes Zoologiske Mu- seet, Copenhagen (UZMC); Ruth Lichtenberg, Naturhistorisches Museum Wien, Vienna (NMW); Christine Dahl, Zoological Institute, University of Uppsala; and L. L. Pechuman, Cornell University, Ithaca (UCI), for the loan of material in their care. We also thank Arnold S. Menke, Raymond Gagne, and Ronald Hodges, Systematic Entomology Laboratory, USDA, Washington; R. M. Reeves, University of New Hampshire, Durham; G. B. Fairchild, University of Florida, Gainesville; and L. L. Pechuman, Cornell University, Ithaca, for their critical review of this manuscript. LITERATURE CITED Austen, E. E. 1908. On the synonymy and systematic position of some species of Tabanidae described by Thunberg and Lichtenstein. Ann. Mag. Nat. Hist. (8) 6: 344-346. . 1922a. Some Siamese Tabanidae. Bull. Entomol. Res. 12: 431-455. . 1922b. Note on a small collection of Indian Tabanidae and other blood-sucking Dip- tera. Trans. R. Soc. Trop. Med. Hyg. 15: 264. Bezzi, M. 1908. Nomenklatorisches iiber Dipteren. III. Wien. Entomol. Ztg. 27: 74-84. Bigot, J. M. F. 1891. Catalog of the Diptera of the Oriental Region. Part I. J. Asiatic Soc. Bengal 60(3): 250-282. . 1892. Description de Dipteres nouveaux (I). Mem. Soc. Zool. Fr. 5: 602-691. Burton, J. J. S. 1978. Tabanini of Thailand above the Isthmus of Kra (Diptera: Tabanidae). Entomol. Reprint Specialists, Los Angeles. 165 pp. Chvala, M. and L. Lyneborg. 1970a. Tabanidae (Diptera) from the Philippines, collected by the Noona Dan Expedition 1961. J. Med. Entomol. 7: 362-366. . 1970b. A revision of Palaearctic Tabanidae (Diptera) described by J. C. Fabricius. 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Philipp. J. Sci. 34: 115-141. Kertesz, K. 1900. Catalogus Tabanidarum orbis Terrarum Universi. Mus. Natl. Hungary, Budapestini (=Budapest). 79 pp. . 1908. Catalogus Dipterorum hucusque descriptorum. Vol. 3. Budapestini. ii + 366 + i pp. Krober, O. 1924. Die Tabaniden der Philippinen. Arch. Naturgesch. 90: 1-27. Lichstenstein, A. A. H. 1796. Catalogus musei Zoologici ditissimi Hamburg!, d II. Februar 1796. Auctionis lege distrahendi. Sectio Tertia Continens Insecta. Verzeichniss von hochstseltenen, aus alien Welttheilen mit vieler Miihe und Kosten zusammen gebrach- ten, auch aus unterschiedlichen Cabinettern, Sammlungen und Auctionen ausgehobenen Naturalien welche von einem Liebhaber, als Mitglied der Batavischen und verschiedener anderer Naturforschenden Gesellschaften gesammlet worden. Dritter Abschnitt, bes- tehend in wohlerhaltenen, mehrentheils auslandischen und hochstseltenen Insecten, die theils einzeln, theils mehrere zusammen in Schachteln festgesteckt sind, und welche am Mittewochen, den 3len Februar 1796 und den folgenden Tagen auf dem Eimbeckschen Haufe offentlich verkauft werden sollen durch dem Mackler Peter Hinrich Packischef- sky. Gottl. Friedr. Schniebess, Hamburg, xii + 222 pp. (The full citation to this work is given as the publication is so rare, and no bibliographies give it in its complete form. According to all published sources, only two copies of this work exist; however, a third copy is present in the University of North Carolina Library). Mitzmain, M. B. 1913. The biology of Tabanus striatus Fabricius, the horsefly of the Phil- ippines. Philipp. J. Sci. 8: 197-221. Moucha, J. 1976. Horse-flies (Diptera: Tabanidae) of the World. Synoptic Catalog. Acta Ento- mol. Mus. Natl. Pragae, Suppl. 7, 319 pp. Nieschulz, O. 1926a. Overbrengingsproeven mit Tabanus rubidus, Wied., T. striatus, Fabr., en Stomoxys calcitrans, L. Dept. Landbouw, Veeartsenijk. Meded., Buitenzorg (=Bo- gor)55. . 1926b. Over de ontwikkeling van Tabanus striatus Fabr. Dept. Landbouw, Veeart- senijk. Meded., Buitenzorg (=Bogor) 56. . 1926c. Over Tabaniden-broedplaatsen op Java en Sumatra. Dept. Landbouw, Veeart- senijk. Meded., Buitenzorg (=Bogor) 58. . 1927a. Overbrengingsproeven met Haematopota cingulata (Wied.). Ned.-Ind. Bladen Diergeneesk. en Dierenteelt 38(5). . 1927b. Ueber Schlupfwespen als Parasiten von Tabaniden-eiern auf Java. Centralbl. Bakt. (Ite. Abt.) 72: 399^110. . 1927c. Ueber mehrfache Infektionen durch Tabanus rubidus, Wied. Centralbl. Bakt. (Ite Abt.) 105: 133-136. . 1928. Enkele miltvuuroverbrengingsproeven met Tabaniden, Musciden en Muskieten. Veeartsenik. Meded. 67, 23 pp. . 1929a. Zoologische Beitrage zum Surraproblem. xxi. Weitere Surraiibertragungsver- suche mit Tabanus striatus Fabr. und T. rubidus Wied. Arch. Protistenkd. 65: 78-96. . 1929b. Enkele miltvuuroverbrengingsproeven met Tabaniden, Musciden en Muskie- ten. Veeartsenijk. Meded., Buitenzorg (=Bogor) 67: 1-23. . 1935a. Ueber die Verbreitung von Tabaniden in Niederlandisch-Indien und ihre wirt- schaftliche Bedeutung. Z. Angew. Entomol. 22: 131-142. . 1935b. Uber die Larvenstadien von Tabanus rubidus Wied. und Tabanus striatus Fabr. Z. Parasitenkd. 7: 639-656. . 1936. Die Entwicklungsstadien von Tabanus rubidus Wied, und T. striatus Fabr. Arch. Naturgesch. (N.F.) 5: 230-255. I I VOLUME 83, NUMBER 2 357 Nieschulz, O. and F. C. Kraneveld. 1929. Experimentelle Untersuchungen iiber die Ueber- tragung der Buffelseuche durch Insekten. Zentralbl. Bakteriol. (1) 113: 403-417. Nieschulz, O. and S. A. S. Ponto. 1927. Zoologische bijdragen tot het Surraprobleem. XVIII. Over meervoudige infecties met Tabanus slriatus, Fabr. Veeartsenijk. Meded., Buiten- zorg (=Bogor) 63, 8 pp. Oldroyd, H. 1957. Some Tabanidae (Dipt.) from Ceylon. Verh. 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