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J. Bequaert.
Notes upon Surcouf's Treatment of the Tabanidæ in the Genera Insectorum and Upon Enderlein's Proposed New Classification of this Family.
Psyche 31:24-40, 1924.

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24 Psyche [February
NOTES UPON SURCOUFJS TREATMENT OF THE
TABANIDA IN THE GENERA INSECTORUM
AND UPON ENDERLEIN'S PROPOSED NEW
CLASSIFICATION OF THIS FAMILY.1
The Tabanidse are a family of Diptera of considerable economic importance. They are universal in their distribution and extremely numerous in species, over 2,100 forms having been described thus far. The great majority of these bite and suck the blood of vertebrates and thus become at times very troublesome to man and his domestic animals, in addition to being actual or potential carriers of infectious diseases. Sur- coufJs review of the family in Wytsman's "Genera Insectorurn" (Brussels, 1921, 205 pp., 5 Pis.) must therefore be greeted with satisfaction. Only those who have attempted work along similar lines can f-ully appreciate the amount of painstaking drudgery and first-hand knowledge involved to make such compilat.ions of real value. Considered as a whole, Surcouf's revision is as satisfactory as it could have been made within a reasonable limit of time and it is far from my intention to present herewith unfavorable criticism. My remarks are merely prompted by the ever increasing interest these flies are assuming for medical and veterinary entomology, so that Surcouf's work is likely to be perused as a source of information by many students with little or no entomological training. It seems therefore necessary to call attention to certain omissions and errors which might easily lead astray the non-specialist.
In the introductory part Surcouf deals with the external morphology and adds certain details of internal anatomy: his researches upon t<he structure of the ocelli and the genitalia are presented as original work. An account of the habits of the adults, oviposition, larval and pupal stages, and enemies follows. This is supplemented by some original observations in an ap- pendix (pp, 186-194) and also by notes under the several genera. Contribution from the Department of Tropical Medicine of Harvard University Medical School.
Pswht 31:24-40 ( 1024). hltp Ilpq~hc enlclub orgrtI/il-024 html



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19241 Notes Upon Surcoufs Treatment af the Tabanidce 25 Nevertheless the treatment of the bionomics is very inadequate and hardly does justice to our present knowledge. Thus it is stated that "the habits of Goniops are unknown" (p. 105)) al- though. the life-history of that genus has been worked out by W. R. Walt,on (Ent. News, XIX, 1908, pp. 464-465, PI. XXII) and W. L. McAtee (Proc. Ent. Soc. Washington, XIII, 1911, pp. 21-29, Pis. 1-111). Incidentally it may be mentioned that Surcouf's supposition that Goniops lives as an external parasite "after the fashion of Hippobosca" is a mere surmise not backed by any observation and highly improbable. To ret,urn to the bionomics of the family, W. Marchand has fortunately published a recent and very full account of "The Early Stages of Tabanidse" (Monogr. of the Rockefeller Institute, New York, No. 13, 1920, 204 pp., 15 Pis.), in which the student will find all needed infor- mation. In his discussion of the parasitary specificity of tabanids (pp. 189-190)) Surc,ouf mentions that, while most of the blood- sucking species attack mammals, Tabanus crocodilinus Austen and other African forms bite crocodiles and that he has himself taken a Tabanus in the Sahara on Varanus griseus. Still more
remarkable, however, is the behavior of Tabanus albipectus Bigot, which, according to Fryer's observations in the Seychelles (Austen, Bull. Ent. Research, XI, 1920, p. 45)) attacks sea- turtles, biting them between t,he plates of the neck. It would be fastidious to list the errors of dates and pages which I have noticed in the bibliography, but the student should be warned against trusting the references indiscriminately. REMARKS UPON THE GENEFL4
Surcouf is extremely conservative in his taxonomic treat- ment, since, with few exceptions, he accepts only genera that have been in use for a long time among students of the group. He retains the division into two subfamilies, Tabaninse and Pangoniinse, proposed more than fifty years ago by H. Lcew (Die Dipteren-Fauna Sudafrika's, I, 1860, pp. 14 and 31). Thanmastocera Grunberg he places at the end of bhe family as a genus of doubtful affinities, but, on account of the absence of tibia1 spurs, it certainly comes in the Tabaninse, a group which,



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26 Psyche [~ebruar~
moreover, contains several other forms with well-developed ocelli.
Surcouf admits 43 genera, of which Baikalia (p. 39; monotypic for B. vaillanti Surcouf), Guyona (p. 141; monotypic for Pangonia mesem brinoides Surcouf, 1908),1 Brodenia (p. 160; monotypic for B. cinerea Surcouf), and Lesneus (p. 161; mono- typic for L. canescens Surcouf) are proposed for the first time. Unfortunately two of these new generic names are preoccupied: Baikalia Surcouf (not Baicalia v. Martens, 1876) I propose to replace by Surcoufiella, new name, and Brodenia Surcouf (not Brodenia Gedoelst, 1913) by Braunsiomyia, new name. The
only species of the last-named genus, Braunsiomyia cinerea (Surcouf) was discovered on the sandy beach at Port Elizabeth (Algoa Bay), Cape Colony, by that enth~siast~ic South African entomologist Dr. H. Brauns.
Walker's subdivisions of Pangonius and most of Ad. Lutz' generic creations among South American t,abanids are not ac- cepted by Surcouf and many of those proposed by Ad. Lutz are not even enumerated. There are, however, a number of other generic names published previous to 1920, which have been over- looked by Surcouf and in some other cases t,he names he uses are obsolete or wrongly spelled.
Hexatoma Meigen, 1820 (p. 26). This name is preoccupied by Hexatoma Latreille, 1809, and should be replaced by Heptatoma Meigen, 1803, which, moreover, has many years priority. Chrysozona Meigen, 1800 (p. 28). I cannot agree with those who claim that this name should replace Hcematopota Meigen, 1803.
I have recently examined an original copy, now at the Library of the American Museum of Natural History, of Meigen's pamphlet "Nouvelle Classification des Insectes Dipthres" (Paris, 1800) and find that this work merely gives short generic des- criptions without mentioning any species, so that these so-called genera having no genotypes should be regarded as nomina nuda, and therefore without nomenclatorial standing. Lepidoselaga Osten Sacken, 1876 (p. 43). There is no suf- ficient reason why this amended form should be preferred to the original Lepiselaga Macquart, 1838.
IGuyona does not appear to be generically distinct from Orgizomyia, as will be shown in a subsequent paper.




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19241 Notes Upon Surcouf's Treatment of the Tabanidoe 27 Dorcalcemus Austen, 19 10 (p. 112). This name was originally spelled Dorcalxmus.
Coenoprosopon Ricardo, 1915 (p. 132). The original spelling of this name is Ccenoprosopon.
Diclisa (p. 112) as characterized by Surcouf is not Diclisa Schiner, 1867, which has as genotype Pangonia incompleta Macquart and is evidently a synonym of Scione Walker, 1850. Surcouf's genus Diclisa appears to correspond to Enderlein's (1922) Rhinotriclista and Triclista.
Cadicera Macquart, 1855 (p. 106).
As shown by Austen
(Bull. Ent. Research, XI, 2, 1920, p. 140)) this name should be replaced by the earlier Phara Walker, 1850. Diatomineura Rondani, 1863 (p. 129). Brethes (Bull. Soc. Ent. France, 1914, p. 59) and Austen (Bull. Ent. Research, XI, 2, 1920, p. 139) have shown that this is a synonym of the earlier Osca Walker (Insecta Saundersiana, Dipt., 1, 1850, p. 10). Orgyzomyia Grunberg, 1906 (p. 139).
The correct spelling
is Orgizomyia.
is name was
Pelecorrhynchus Macquart, 1850 (p. 110). Th' originally spelled Pelecorhynchus.
The following generic names are not listed by Surcouf: Acanthocerella Brhthes, An. Mus. Nac. Buenos Aires, XX, 1910, p. 475. Monotypic for A. boliviensis Brhthes, 1910. South America.
Arnphichlorops Ad. Lutz, Mem. Inst'. Osw. Cruz, VI, 1914, p. 166. Type : Tabanus flavus Wiedemann, 1828. South America. Anacampta Schiner, Verb. Zool. Bot. Ges. Wien, XVII, 1867, p. 305. Without description or species. Evidently an error for Apocampta Schiner.
Catachlorops Ad. Lutz, Mem. Inst. Osw. Cruz, VI, 1914, p. 166. Type : Dichelacera fuscipnnis Macquart, 1847. South America.
Chelotabanus Ad. Lutz, Mem. Inst,. Osw. Cruz, VI,1914, p. 166. Type: Tabanus fuscus Wiedemann, 1819. South America.
Chlorotabanus Ad. Lutz, Mem. Inst. Osw. 'Cruz, VI, 1914, p. 167. Type: Tabanus mexicanus Linnaeus, 1767. North and South America.




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28 Psyche [~ebruar~
Cosnura Bigot, Ann. Soc. Ent. France, (3) V, 1857, p. 286. Monotypic for C. longicauda Bigot,, 1857. Chile. A number of species have been described, all of which have been omitted by Surcouf.
Cryptotylus Ad. Lutz, Mem. Inst. Osw. Cruz, VI, 1914, p. 166. No species mentioned. South America. Cydistomyia Taylor, Proc. Linn. Soc. New South Wales, XLIV, 1919. p. 47. Monotypic for C. doddi Taylor, 1919. Queensland.
Dieladocera Ad. Lutz, Comm. Linhas Telegr. Estr. de Matto Grosso ao Amazonas, Ann. No. 8, Zool., Taban., 1912, p. 4. Monotypic for Dicladocera unicolor Ad. Lutz, 1912. Dyspanqonia Ad. Lutz, Revista Soc. Scientif. Sao Paulo, I, 1, 1905, p. 27. Type: Pangonia fuscipennis Wiedemann, 1828. This is a synonym of Esenbeckia Rondani. Erephosis Bigot, M6m. Soc. 2001. France, IV, 1891, p. 414. Misspelling of Erephopsis.
Esenbackia Surcouf, Bull. Mus. Hist. Nat. Paris, XV, 1909, p. 257. Misspelling of Esenbeckia.
Gonisops Kertesz, Catal. Tabanid., 1900, p. 25. Evidently
a misspelling of Goniops.
Holococeria Ricardo, Arch. f. Naturgesch., LXXX, Abt. A, Heft 8, (1914) 1915, p. 128. A misspelling of Holcoceria. Laphriopsis Ad. Lutz, Mem. Inst. Osw. Cruz, 111, 1911, p. 71. An evident error for Laphriomyia. Leptotabanus Ad. Lutz and A. Neiva, Mem. Inst. OSW. Cruz, VI, 1914, p. 72. The name is used in an enumeration of species for Leptotabanus nigrovenos& Ad. Lutz and A. Neiva, but I was unable to find a description of either genus or species. Leucotabanus Ad. Lutz, Mem. Inst. Osw. Cruz, VI, 1914, p. 167. Type: Tabanus leucaspis Wiedemann, 1828. South America.
Macrocormus Ad. Lutz, Mern. Inst. Osw. Cruz, VI, 1914, p. 167. Type: Tabanus sorbillans Wiedemann, 1828. South America.
Melanotabanus Ad. Lutz and A. Neiva, Mem. Inst. OSW. Cruz, VI, 1914, p. 76. Monotypic for M. fuliqinosus Ad. Lutz and A. Neiva, 1914. Sout,h America.




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19.241 Notes Upon Surcoufs Treatment of the Tabanidoe 29 Metoponaplos Ricardo, Arch. f. Naturgesch., LXXX, Abt. A, Heft 8, (1914) 1915, p. 124. Type by original designation: Pangonia parva Walker, 1848. According to Enderlein (Mitt. 2001. Mus. Berlin, X, 2, 1922, p. 342)) this is a synonym of Scarphia Walker, 1850, a view with which I fully concur. Merycomyia Hine, Ohio Naturalist, XII, 1912, p. 515. Type by original designation: Tabanus whitneyi Johnson, 1904 (Syn. : Merycomyia geminata Hine, 1912). North America. Neochrysops Walton, Proc. Ent. Soc. Washington, XX, 1918, p. 191. Monotypic for N. globosus Walton, 1918. North America.
Neotabanus Ad. Lutz, Mem. Inst. Osw. Cruz, VI, 1914, p. 167. Type : Ta banus trilineatus Latreille, 1814. South America. Ad. Lutz (Ibidem, p. 47) claims that his generic name has priority over Neotabanus Ricardo, 1911, but I have been unable to discover on what evidence this statement is based. Orthostylus Ad. Lutz and A. Neiva, Mem. Inst. Osw. Cruz, VI, 1914, p. 74. Monotypic for 0. ambiguus Ad. Lutz and A. Neiva, 1914. South America.
Palimmecomyia Taylor, Proc. Linn. Soc. New South Wales, XLII, 1917, p. 518. Monotypic for P. celcenospila Taylor, 19 17. Queensland.
Parasilvius Ferguson, Proc. Roy. Soc. Victoria, N. S., XXXIII, 1921, p. 8. Monotypic for P. fulvus Ferguson, 1921. Aust,rali a.
Phceomyia Ad. Lutz, Mem. Inst. Osw. Cruz, 111, 1911, p. 83. Evidently a misspelling for Phceoneura. Phceotabanus Ad. Lutz, Mem. Inst. Osw. Cruz, VI, 1914, p. 168. Type : Tabanus litigiosus Walker, 1850. South America. Phibalomyia Taylor, Proc. Roy. Soc. Victoria, N. S., XXXII, 2, 1920, p. 165. New name for Elaphromyia Taylor (Proc. Linn. Soc. New South Wales, XLI, 1917, p. 749), not of Bigot, 1859. Monotypic for Elaphrom yia carteri Taylor, 19 17. Queensland. Philorites Cockerell: Entomologist, XLI, 1908, p. 264. Monotypic for P. johannseni Cockerell, 1908. Fossil in the Eocene of Colorado.




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30 Psyche [~ebruar~
Rhabdotylus Ad. Lutz, Mem. Inst. Osw. Cruz, VI, 1914, p. 166.
Type : Tabanus planiventris Wiedemann, 1828. South America.
Poecilosoma Ad. Lutz, Mem. Inst. Osw. Cruz, VI, 1914, p. 167. Type: Tabanus quadripunctatus Fabricius, 1805. South America.
Rhigioglossa Wiedemann, Aussereurop. Zweifl. Ins., I, 1828, p. 105. Used in the combinaction "Rhigioglossa testacean as a synonym of Rhinomyza edentula Wiedemann, which thus will be its genotype. The name takes precedence over Erodior- hynchus Macquart, 1838, based upon the same species. Rhynomyza Surcouf, Bull. Mus. Hist. Nat. Pazis, XV, 1909, p. 260. Misspelling of Rhinomyza.
Stenotabanus Ad. Lutz, Mem. Inst. Osw. Cruz, VI, 1914, p. 167. Type: Tabanus tmiotes Wiedemann, 1828. South America
Stictotabanus Ad. Lutz and A. Neiva, Mem. Inst. Osw. Cruz, VI, 1914, p. 72. The name is used in an enumeration of species for Stictotabanus maculipennis (Macquart) . Since this is a described species the generic name has a standin,. o- in a nomen- clature, even though the genus has apparently not been hitherto defined.
In the case of the new generic names proposed by Ad. Lutz in Mem. Inst. Osw. Cruz, VI, 1914, pp. 166-168, the only pub- lished descriptions are contained in a key and are not accom- panied by references to species. In a previous article by Lutz and Neiva, however, which appeared in t'he same volunie (pp. 69-80), these new names have been used in enumerations of species and I have selected genotypes from among them. Cryp- totylus alone has apparently not yet been used in connection with a specific name so that it still is a nomen nudum. Ad. Lutz also closes his article with the statement (p. 168) that it was published before (in 1913) in the Brazilian journal "Brazil Medico." I have been unable to discover whether his new generic names should therefore be properly dated from 1913, but it would appear that they were not used in connection with specific names previously to 1914. Many of Ad. Lutz' proposed genera have not been noticed in the Zoological Record.



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19241 Notes Upon Surcouf's Treatment of the Tabanidoe 31 The following additional genera of Tabanidse are of more recent date:
Heterochrysops Krober, Zool. Jahrb., Abt. f. Syst., XLIII, 1920, p. 55.
For a number of Palaearctic species of Chrysops, none of which is designated as type. Chrysops flavipes Meigen, 1804, is herewith selectled as such.
Neochrysops Szilhdy, Ann. Hist. Nat. Mus. Hungarici, XIX, 1922, p. 126. Type by original designation: Neochrysops grandis Szilhdy, 1922, Formosa. The name is preoccupied by Neochrysops Walton, 1918.1 The genus, however, appears doubtfully distinct from Chrysops and need therefore not be renamed at present.
Silviochrysops Szilhdy, Ann. Hist. Nat. Mus. Hungarici, XIX, 1922, p. 126. Monotypic for Silviochrysops flavescens Szilhdy, 1922, Formosa.
Surcoufia Krober, Arch. f. Naturgesch., LXXXVIII, Abt. A, Heft 8, 1922, p. 115. Monotypic for Surcoufia paradoxa Krober, 1922, Northwest Africa.
Finally in a recent paper which will be considered in detail below; Enderlein has proposed a considerable number of new generic names. These it appears unnecessary to list at present, since their exact status is a? yet uncertain. Among the misspellings of names, I mention only those of Chr ysops nigribimbo Whitney (not niqrilimbo) and Tabanus superjumentarius Whitney (not sube-rjumentarius). Tabanus lugubris Osten Sacken appears to belong properly in the genus Snowiellus, from examination of a specimen obtained at Tampa, Florida, by Mr. E. Bell.
Tabanus whitneyi Johnson belongs in the genus Merycomyia. Tabanus mexicawus. The synonymy and distribution given by Surcouf under that name should be revised. As shown by I?. Knab (Insecutor Inscitise Menstruus, IV, 1916, pp. 95-loo), four species have been commonly confused under mexicamis: Weochrysops Bethune-Baker, Trans. Ent. Soc. London, (1922) 1923, p. 279, in Lepidoptera, is similarly preoccupied.



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32 Psyche [~ebruar~
(1) Tabanus mexicanus Linnaeus (Syn.: T. olivaceus de Geer and T. punctatzis Fabricius).
Mexico, Central America,
Trinidad.
(2)
Tabanus inanis Fabricius (Syn. : T. ochroleucus Meigen, 5". viridiflaws Walker, and perhaps also T. sulphureus Palisot de Beauvois). South and Central America.
(3) Tabanus flavus Macquart . Southeastern United States : from New Jersey to Florida, Missouri and Louisiana. This name is unfortunately preoccupied by Tabanus flavus Wiede- mann and no substitute appears to be available. (4)
Tabanus luteoflaz~us Bellardi (Syn.: T. mexicanus var. limonus Townsend). Mexico.
The following North American species have been omitted: Chrysops calopterzis Hine, Ohio Naturalist, VI, 1905,~. 392, Q . Guatemala.
Chrysops hinei Daecke, Ent. News, XVIII, 1907, p. 143, 9. New Jersey.
Chrysops paruulus Daecke, Ent. News, XVIII, 1907, p. 142, 9. New Jersey.
Merycomyia qeminata Hine, Ohio Naturalist, XII, '1912, p. 515, PI. XXV, figs. 2 and 4, 9 8'.
This is a synonym of
Merycomyia whitneyi (Johnson).
Merycomyia mixta Hine, Ohio Naturalist, XII, 1912, p. 516, PI. XXV, figs. 1 and 3; Q . Georgia, North America. Neochrysops
Walton, Proe. Ent . Soc. Washington,
XX, 1918, p. 192, fig. f, 9 . Eastern United States (Maryland), Silvius jonesi Cresson, Proc. Ac. Nat. Sc. Philadelphia, LXXI, 1919, p. 175, Q 8'.
California.
Tabanus subniqer Coquillett, Ent. News, XVII, 1906, p.
48, Q . Illinois.
Tabanus atratus Tar. fulvopilosus Johnson, Psyche, XXVI, 1919, p. 164, 9. Florida, New Jersey.
A number of addit,ional species of Tabanus have been des- cribed by J. McDunnough in 1921 and 1922 (Canad. Entom., LIII, 1921, pp. 139-144 and LIV, 1922, p. 239)) namely: T. atrohis, T. laniferus, T. metabolus, T. nudus, T. rupestris, and T. trepidus; and this author has also reinstated Tabanus calif-



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19241 Notes Upon Surcouf's Treatment oj the Tabanidce 33 ornicus Marten and T. hcemaphorus Marten as valid species. More recently Hine (Canad. Entom., LV, 1923, pp. 143-146) has added T. gracilipalpis and T. sexfasciatus. Including these, we obtain a total of 334 species of Tabanidse known at present from America north of Panama. They are divided among the several genera as follows: Apatolestes, 1; Chrysops, 7 1 ; Hcematopota, 3 ; Corizoneura, 4; Diachlorus, 1 ; Diatomineura, 4; Dichelacera, 6; Rhinotriclista (Diclisa of Sur- couf) , 1 ; Erephopsis, 2 ; Esenbeckia, 1 ; Goniops, 1 ; Lepiselaga, 1 ; Merycomyia, 2; Neochrysops, 1 ; "Pangonius,^1 18; Pityocera, 1; Scione, 2; Silvius, 4; Snowiellus, 2; Stibasoma, 2; and Ta- banus, 206. Of these Goniops, Merycomyia, Neochrysops, and Snowiellus are restricted to the Nearctic region (north of Mexico). It may still be mentioned that Surcouf (p. 130) erroneously quotes Trichophthalma amcena Bigot and Hermoneura landbecki Philippi among the synonyms of Diatomineura latipalpis (Macquart), having evidently followed in this Kertesz (Cat. Dipt., 111, 1908, p. 170). Both Bigot's and Philippi's descrip- tions refer, however, to a nemestrinid which should be known as Eurygastromyia landbecki (Philippi). See Lichtwardt, Deutsch. Ent. Zeitschr., 1910, p. 608.
Tabanus corax Law, Wien. Ent. Monatschr., VII, 1863, p. 10.
Surcouf (p. 79) lists this as a doubtful synonym of Ta- banus pluto Walker. Neave (Bull. Ent. Research, V, 1915, p. 308)) however, has shown that Lcew's name should be used for Tabanus xanthomelas Austen, of which T. leucaspis v. d. Wulp (not of Wiedemann) is a synonym.
. Tabanus alboventralis Newstead is recorded twice in the list (p. 59), the first time misspelled "albiventralis." It is ap-
parently a synonym of T. sufis Jaennicke. lWhether there any true Pangonius, in the restricted sense, in North America appears extremely doubtful.
The three species which I have ex-
amined, viz., tranquilla Osten Sacken, rasa Osten Sacken, and fora Williston present all the characters of Austen's genus Buplex, to which, I believe, they should be transferred.




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34 Psyche [February
Tabanus blanchardi Surcouf and T. gabonensis Macquart are still listed as distinct species, whereas they are now generally regarded as synonyms of T. secedens Walker. Hcematopota maculosifacies Austen is listed twice (p. 34) ; also erroneously as maculifacies Austen. Pangonius brevis Austen (p. 127) belongs properly in the genus Phara (Cadicera) ,
Pangonius austeni J. Bequaert (p. 127). This name should be deleted from the list. As stated by Austen (Ann. Mag. Nat. Hist., (8) XI, 1913, pp. 560-562)) it was based upon the male of P. infuscus Austen and the female of Diutomineura neavei Austen. I have convinced myself of the correctness of Major Austen's view after he has kindly compared some of my specimens with the types of these two species, during my recent visit at the British Museum.
Pangonius neavei J. Bequaert (p. 128) should also be dropped since it is not Austen's Diutomineura neavei, but was based on both sexes of Corizoneura inornate, Austen, as I have recognized after comparison with the types.
In my paper on Congo tabanids, Rev. Zool. Afr., 11, 19133 p. 222, I have also recorded a male Chrysops fusca and a female C. distinctipennis. As Major Austen has pointed out to me at the British Museum, both specimens belong to Chrysops stig- maticalis Lcew. On the other hand, Diutomineura virgata Austen, DorcaZoemus candidolimbatus Austen, and D. compactus Austen of the same paper were correctly identified. I have noticed the following misspellings of names: Hcema- topota heptogramma for H. heptagramma; H. hirsutitursis for H. hirsutitarsus; Tabanus nyassce for T. nyasce; and T. wosnami for T. woosnami.
The following Ethiopian species have been omitted: Hcematopota furva Austen, Bull. Ent. Research, 111, 19 12, p. 334, PI. XI, fig. 7, Q 8. Uganda and Kenya Colony. Hcematopota pertinens Austen, Ann. Mag. Hat. Hist., (8)


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