J. Bequaert.
Notes on Hippoboscidae. 13. A Second Revision of the Hippoboscinae.
Psyche 46(2-3):70-90, 1939.

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Psyche
[ June-Sept.
NOTES ON HIPPOBOSCID^E
13. A SECOND REVISION OF THE HIPPOBOSCINAE Harvard Medical School and School of Public Health, Boston, Mass.
Since the publication of my synopsis of the Hippoboscinse (1931, Psyche, XXXVII, (1930), pp. 303-326), much addi- tional information has come to light. O'ne new species was described recently by the late G. A. H. Bedford and I was able to study several types, including those of H. fulva Austen.
In the alphabetical list of names (p. 306), martinaglia Bedford should be inserted as the ninth valid species; longi- pennis Fabricius is the valid name of capensis v. Olfers, which becomes a synonym; and varieguta Megerlk (not to be credited to Wiedemann) is the valid name of maculuta Leach, which passes in the synonymy.
As pointed out before, the Hippoboscinse differ from other members of the family in several important charac- ters. To those listed before may be added the presence of a pair of deep, more or less pit-like depressions, placed later- ally on the suture between mesonotum and scutellum; also the well-defined pale yellow or white spots of head and thorax, which are not duplicated elsewhere in the family. While in other Hippoboscidse color differences are of little or no specific value, in Hippobosca the shape and arrange- ment of the pale spots produces a pattern to a large degree diagnostic for each species. In this respect, there is an ob- vious analogy to the characteristic pattern of pale spots found in many species of ticks (Amblyomma and Derma- centor).
I have been at pains to discover additional specific charac- ters, particularly in the case of closely allied forms. The chetotaxy has been neglected thus far, yet offers reliable dif- ferences which should be investigated by the accepted bio- Pzirfif W70-90 (1939). hup //psyche rnlclub omfW-W-070 html



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19391 Notes on Hippoboscidae 71
metrical methods.
My material is not extensive enough
for the purpose.
Considering only the chetotaxy of the
scutellum, this sclerite in Hippobosca bears at the extreme apical margin and somewhat ventrally a dense fringe of short, soft hairs. Anterior to the fringe one finds groups or rows of long, stiff bristles, either black or pale-colored, which I shall call the preapical bristles. The groups may be either far apart and restricted to the extreme sides or more or less connected medially. In some species the bristles are placed in one row, in others they form two irregular rows or are merely bunched together. When there are many bristles, these are often mixed with a few soft, short hairs, sometimes forming a second row behind the stiff bristles. In most cases the number and arrangement of the preapical bristles is the same in both sexes. There are two exceptions. In H. struthionis the males have more bristles on the aver- age than the females. In the males of H. camelinu the pre- apical bristles occupy the same position as in the other spe- cies of the genus; but in the females, the bristles are placed nearer the middle, being rather discal or medio-scutellar, and are also fewer in number than in the males. Except for this case of the female H. camelina, there are no discal nor basal bristles, setae or hairs on the scutellum in Hippobosca. The variation of the preapical bristles of the scutellum is discussed under each species, but the following summary compares the species for diagnostic purposes. The number of specimens examined is given in parenthesis. equina (106) : 5 to 11 bristles Q 8
(average, 7).
longipennis (143) : 3 to 7 bristles $ 8
(average, 5).
fulva (3) : 8 bristles $ 8 .
vuriegutu (95) : 13 to 27 bristles $ 8 (average, 18). rufipes (95) : 12 to 23 bristles 8 (average, 17). hirsuta (6) : 14 to 18 bristles
8.
martinaglia (not seen).
struthionis (23) : 4 to 12 b.
$
(average, 6) ; 8 to 15 b. $
(average, 10).
camelina (31) : 3 to 8 b. $ (average, 5) ; 11 to 22 b. 8 (average, 14).
In Hippobosca, the integument of the abdomen, behind the usual large tergal and small sternal sclerotized basal plates, is mostly soft and extensible. All species I have seen have



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72 Psyche [ June-Sept.
two pairs of sclerotized subapical (lateral) plates in both sexes. In the males of equina and longipennis, the anterior pair is small and fused with the median tergal plate, yet recognized by the very long setae it bears. In addition, these two species have in both sexes three median tergal sclerotized plates, which are much smaller in the female than in the male. Median tergal plates are lacking in both sexes of variegata, mfipes, struthionis, and hirsuta, and in the female of mmelina.. The male of camdim, however, has an extensive anterior sclerotized median plate, immediately behind the basal tergal sclerite of the abdomen, and poste- riorly a pair of small, median tergal plates. In the female of longipewlis and equina the anterior pair of subapical (lateral) plates is slightly smaller than the posterior pair ; it is larger than the posterior pair in the female of vanag~ta, hirsuta, camelina and struthionis; and both pairs are about the same size in the female of rufives. In the males of vaviegata, mfipes, hirsute,, cumelim and strut hionis, the anterior pair of subapical plates is very large, the posterior pair very small and readily overlooked. I have not exam- ined the structure of the abdomen of fdva and martinaglia. In the male genitalia, the claspers (or parameres) are very similar in equina,, longipennis, fwiva, variegata, rufipes and struthionis, being more or less slender, straight and rod-like, ending in a point.
In camelha they are of much the same
rod-like type, but thicker, curved in profile and ending in a blunt, somewhat knobbed point. They are quite aberrant in hirsuta, being thick and beam-like, with a broadly trun- cate and slightly emarginate tip. The genitalia of martin- ugh are unknown.
To sum up, the nine species of Hippobosm now recognized may be divided into four groups, expressing relationship based on structural characters ; (1) equina, longipennis and fulvu; (2) variegata, ntfipes, hirsuta and possibly martin- a&; (3) stmthiomis; (4) camelina,.
The following key supersedes that of my earlier paper (pp. 308-309). H. martinagdia is inserted from the descrip- tion only.
1. Second longitudinal vein (Ra + 3) long, about as long as or longer than last section of third longitudinal, reaching costa much beyond tip of first longitudinal



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19391 Notes on Hippo boscidae 73
(Ri) and usually apicad of anterior cross-vein (r-m) ; last section of costa about three times the length of penultimate section or shorter. Base of third longi- tudinal vein (R4 + 5) bare. One pair of vertical bristles. Preapical bristles of scutellum few (3 to 11). Abdomen with three median tergal sclerotized plates in both sexes (in equina and longipennis; not known in fulva).
Two pad-like pulvilli at sides of
bristle-like empodium, one much larger than the other. Parameres of male genitalia slender, rod-like, ........................................................ ending in a point 2
Second longitudinal vein short, shorter than last section of third longitudinal, reaching costa together with or close to tip of first longitudinal; last section of costa at least five times the length of penultimate section.. .4 2. Larger species, the wing 6 to 8.5 mm. long. Apical
lobes of fronto-clypeus irregularly and broadly tri- angular, their inner margins curved. Scutellum f uscous to f erruginous laterally, yellowish-white medially, rarely more extensively yellowish ; with a regular row of 5 to 11 preapical bristles (usually 6 to 8), divided into two groups. Wing veins as a rule rufous to dark brown ............................ H. equina Smaller, the wing at most 6 mm. long. Scutellum as a rule entirely or nearly entirely yellowish or ivory- white .............................................................................. 3 3.
Wing 5 to 6 mm. long. Apical lobes of fronto-clypeus regularly and sharply triangular, separated by a broad notch, their inner margins nearly straight. Scutellum with 3 to 7 preapical bristles (usually 5 or 6). Wing veins mostly pale testaceous, usually with some darker stretches ................ H. longipennis. Wing 4.2 to 4.5 mm. long. Apical lobes of fronto-clypeus irregularly and broadly lobular, separated by a nar- row slit, their inner margins curved. Scutellum with about 8 preapical bristles ................................ H. fulva 4. Base of third longitudinal vein (R4+5) setulose over some length on the upper side. One pair of vertical bristles. No median tergal plates in both sexes (in variegata, rufipes and hirsuta; probably also in mar-



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74 Psyche [ June-Sept.
tinagliu) . Only one pulvillus well-developed, the other rudimentary ...................................................... 5 Base of third longitudinal vein bare .............................. 8
5. Second longitudinal vein very short, forming an oblique cross-vein which ends in the first longitudinal and runs from opposite or apicad of upper tip of anterior basal cro,ssvein (M3) to basad of anterior cross-vein (r-m) . Frons distinctly narrower at occiput than at fronto-clypeus, the postvertex much longer than wide. Scutellum as a rule with three ivory-white spots, the largest in the center; with 13 to 27 preapical bristles (usually 16 to 20), in one or two irregular and fairly continuous rows. Parameres of male genitalia slen- der, rod-like, ending in a sharp point. Wing 7 to 8 mm. long .................................................... H. variegata. Second longitudinal vein longer and more slanting, end- ing in costa at or beyond tip of first longitudinal and running from basad of upper tip of anterior basal cross-vein to opposite or basad of anterior cross-vein ........................................................................................ 6 6.
Smaller, the wing 4.5 mm. long. Frons wide, not ap- preciably narrower at occiput than at fronto-clypeus, the postvertex much wider than long. Mesonotum reddish-brown, with an anterior median dark band extending posteriorly to near the transverse suture where it is more or less forked. Scutellum entirely yellowish-white ; with relatively few (probably eight to ten) preapical bristles. Mesonotum moderately bristly, bare in the center .................... H. ma~tinaglia Larger, the wing 6.5 to 9 mm. long. Frons narrower, the postvertex nearly as long as wide or slightly longer. Color pattern of mesonotum different. Pre- apical bristles of scutellum more numerous (12 to 23) .................................................................................. 7 7.
Frons distinctly narrower at occiput than at fronto- clypeus. Scutellum very wide and nearly rectangular, with a median, rufous and two lateral, ivory-white spots; with 12 to 23 heavy preapical bristles (usually 14 to 20), placed in one regular, almost continuous row. Mesonotum moderately bristly, bare in the cen-



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19391 Notes on Hippoboscidae 75
ter. Legs bright reddish-brown. Parameres of male genitalia slender, rod-like, ending in a blunt point. Wing 7 to 9 mm. long .................................... H. mfipes Frons very slightly or not narrower at occiput than at fronto-clypeus. Scutellum narrower and less rec- tangular, with a median ivory-white spot; with 14 to 18 soft and pale preapical bristles, placed in two irregular, more or less connected groups. Mesonotum very bristly, a1,so in the center. Legs rufous-yellow. Parameres of male genitalia thick, beam-like, trun- cate and slightly emarginate at tip. Wing 6.5 to 8 mm. long ........................................................ H. hirsuta 8. Two or three pairs of vertical bristles. F'ronto-clypeus shorter than its distance from the occipital margin. Postvertex shorter than mediovertex, the latter much narrowed medially by the broad inner orbits. Ante- rior basal cross-vein (M3) very oblique and nearly its own length from anterior cross-vein (r-m) . Scutel- lum semi-elliptical, the hind margin distinctly convex and slightly projecting medially ; in the female with 3 to 8 discal bristles (usually 4 to 6), placed in two linear groups; in the male with 11 to 22 preapical bristles (usually 12 to 15), placed in two irregular groups.
No median tergal plates in the female; the male with one large median plate behind the basal tergal sclerite and a pair of small median tergal plates posteriorly. No pad-like pulvilli ; bristle-like em- podium bare, except at base. Parameres of male genitalia rod-like, but curved upward, the apex slight- ly swollen and knob-like. Wing 9 to 10 mm. long ...................................................................... H. camdina One pair of vertical bristles. F'ronto-clypeus nearly as long as its distance from occipital margin. Post- vertex as long as or longer than mediovertex, the lat- ter moderately narrowed by the inner orbits. Ante- rior basal cross-vein short, almost vertical upon the fourth longitudinal and more than twice its length from anterior cross-vein. Scutellum nearly rectang- ular, the hind margin more straightly truncate; in both sexes with preapical bristles placed in two widely separated lateral groups, more numerous in the male



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(8 to 15, usually 9 t~ 12) than in the female (4 to 12, usually 6 to 7). Abdomen in both sexes without median tergal plates, Two pad-like pu1villi; empo- dium feathered. Parameres of male genitalia den- der? rod-like, ending in a sharp point, Wing '7 to 7.6 mm, long ..
1. Hippobowa quim Linn~us. -
ha die" is near Smyrna.
Additional Specimens Examined. - Norway : Smaalen-
ene. - Esthonia, one female off a duck, Mergus (or Mw- gamer} sermtw Linnaeus (sent by G. B. Thompson), an accidental host. - Finland : Kusa ( C. Lundstrh) . - Denmark : SeelIand (Univ. Zml. Mug., Copenhagen). - Awtria : Grunbach, Schneehrg region (Handlimch) , - Hungary. - Roumania : Eihar Mts. (K. Jordan) ; Hex- kulesbad (W. Rothschild and E. Hart&). -Bulgaria: Aladza near Vaxna; Bela Cerkva, Rhodope (Zexny).- Jugo-Shia : Zlj eb, New Montenegro ; Stdac, Herzegovina {Penther) ; Bosnia. - Frame: Argentat, Auvergne, off a cow. - Italy: T~iest ; Taranto ; Fala. - Corsica : Vima- vona (M. E. Mody) ; Corte (M. 33. Mosely 1 ; La Poce de Vizzavona (Yerbuxy ) ; Ajaccio (F. Gugliemi) . - Spain : Mureia (G. L. Boag) ; Sierra de Guadarxama, 6,000 to 8,000 ft. (B. Uvaxov) ; Algeciras (Zemy) ; Noguera near Albax- xacin? Aragon (Zerny) . - Canary Islands : La CaIdera, Las Palmas Id. (W. M. Wheeler) ; Sa Cmz, Tenerifle; Puerto Cabias, Buenaventuxa. - Madeira (Lowe). - Albania: Kda Ljums ; Hodzha near F~izren ; Fashtrik ; Korab ; Du- ramo. - Greece: Stavros, Macedonia (J. Waterston) ; Saloniki (J. Waterston) ; Helmas (Fonberg) ; Taygetos ; KoystalIopyghi (A. H. G. Alstoni) ; Attica; Poxos ; Vrissula; Mt. Pangaion ; Struma ; Carvalli (R. C. Shannon). - Asia Nfnor (Amtolia}: Mamerun; Cilician Taurus (Prince Abemperg) ; Sabmdscha to Eskischebir ; Ephesus. - Trans- caucasia: Sagah on Lake Goktschai (Zugmayer). - Turke- stan. - Persia {Iran) : Nissa, Elburs Mts. (Brandt) ; Tsiang-Kanspe, E. Persia (A. Teufigi) ; Emeli, N. W. Persia (P. A. Buxtun). -Arabia: Akaba, Hejaz (W, M. Marin). - Palestine: Haifa (P. J. Bamaud) ; Beisan? Jordan Valley (P. A. Buxton) . - Cyprus: Limasol (G. A. Mavromow- takis) . - Egypt: Tisfa {Zool. Depk Univ. Egypt). -



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19391 hrotes on Hippoboscidae 77
Libya : Dernah (Klaptocz) . - Tunis : Gabes (Mik) . - Al- geria : Hamman Rirha ; Hamman Meskoutine ; El Kantara ; Biskra (W. Rothschild and E. Hartert).-Morocco: Aguelman Sidi Ali bu Mohammed, Middle Atlas, 6,500 ft. (K. Chapman and J. W. S. Pringle) ; Ijoukak, Great Atlas, 3,900 ft. (K. H. Chapman and G. A. Bisset) ; Arround, Atlas, &950 m. (Ebner). -Australia:
Sydney, on a horse
imported from New Caledonia. - New Hebrides : Tanna Id. (E. Robertson) ; Vila, Efate (or Sandwich) Id., very com- mon on horses (P. A. Buxton) . - Philippines : Alabang, Rizal (M. B. Mitzmain). - Amboyna, off cattle (F. Muir). - Singapore (F. Muir) .
The preapical bristles of the scutellum vary from 5 to I1 in 106 specimens examined (58 9 and 48 8 ), from 22 10- calities, 85 specimens having from 6 to 8 bristles. The specimens fall in the following groups: with 5 bristles: 3 (I 9,2 8);6b.: 28 (17 Q, I1 8);7b.:22 (9 Q, 13 8); 8b.:35 (24 ?,I1 8);9b.:IO (4 ?,6 8);10b.:7 (3 Q?4 8 ) ; Ilb. : 1 ( 8 ) . There is no evidence of any sexual dif- ference in this character.
No locality is represented by
enough specimens to make a further analysis of any sig- nificance. The bristles are evenly divided between both sides in 69 specimens, unevenly in 37? the uneven groupings observed being 2 + 3, 3 + 4, 3 + 5, 4 5, and 6 + 6. The frons, in the male? is nearly parallel-sided and about as wide as an eye; in the female it is slightly widened me- dially, where it measures a little over the width of an eye. In addition to the average larger number of preapical bristles of the seutellum, H. equina differs also from H. longipennis in the shape of the tergal plates of the abdomen. In the female of H. equina) the three median, setulose ter- gal plates are larger than in the female of Zongipemis, more transverse and ribbon-shaped, the median plate only slight- ly smaller than the hindmost plate? which is nearly as large as the anterior pair of subapical (lateral) plates. The median plates bear many more setae than in longipennis. In the male of equina) the three median tergal plates are large and ribbon-like, the hindmost (or third) plate fused with the anterior pair of ovate? widened subapical (lateral) plates (bearing longer bristles than the median plate proper). The shape and arrangement of the tergal plates



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78 P8gch~ [ June-Sept.
are shown correctly by Ferris (1930, Philippine Jl. Sci,? XLIII? p. 540? fig. I Q , and p, 643, fig. 4a 8 1. The dasper~ (or parameres) of the male geniblia are slender, rod-like and xegdaxly pointed at apex. They are figured in ~ide view by J. I. Roberts (1927, Ann. Trop Med. Paras.? XXI, PI. 111, fig. 8) ; Been from above they are much narrower,
2* Hippobosca hngipmmis Fabricius, 1805. - A study of Fabricius' two types, marked "ex Tranquebar (MUK Dom. Lund) ", at the University Zoological Museum, Copen- hagen? shows that they are the species commonly known a8 H. capewh v. Olfer~ (l8l6), H. frumilloni Leach {lBlT), ox H. wminu Rondani (1878). Why Fabricius wrote "Caput et thorax fermginea immaculata" is a mystery, as both hi8 types show the characteristic capemis pattern. A specimen from Kalewa, Upper Burma? named %ngipemnk" by Major Austen, wag &udkd at the British Museum and showed no structural characters differentkxting it from the usual H. cupen&.
Fabrkius' name antedates all 0th~ designations fox the species, which is extremely variable in color. I have also seen, at the British Museumy a specimen of H. fmn- cilloni, apparentIy labelled by Leach and which may be the type, although it is not marked as such. Addjtional Specimens Examined. - Bulgaria : Sredne near Russe. - Greece: Stmma (R. C. Shannon). - Trans- caucasia: Saga111 on Lake Gokkhai (Zugmayer). - Transca~pia : Imam-Baba, Merv District (L, Mi&schenko). -Persia (Iran) : Keredjv 40 Kilom. from Teheran {Brandt) ; Dschulfa? N, W. Persia. - Mesopotamia (Iraq) : Djerabis; Asmr; Dawah [A. D. Fraser) ; Bagdad (T, C. Connor) ; Amara (T. C. Connor). - Syria : Beirut (E. S. Sewell). -Palestine: Amman, E. of Jordan (F. A Buxton). - Arabia : Ras Farkak, Hadramaut (Simony). - Cypxu8 : Lamaw, off cow.-Egypt: Tisfa (ZooL Dept. Univ. Egypt). - Libya: Bengasi. - Tunis : Gabhs. - Morocco: Marrakeah (G, 3. Fairchild). - Anglo-Egyptian Sudan: Lugnd, Darfur Province (H. Lynes) ; Khor Arbaat, Fort Sudan, - Kenya Colony : Lake Jipe ; Marsabity Rerdili Nymo (C. A. Neave) ; Turkana District; Mexifano (Mc- Arthur) ; Voi (Tab) ; Lemek, Masai Reserve (A. 0. Luek- man). - Tanganyika Territory: Valley of Ruaha River? N. Uhehe; Morogom, Uluguru (A. G. Wilkins) ; W. shore of



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19391 Notes on Hippoboscidae 79
Lake Manyara (B. Cooper) ; Mt. Meru? off lion (B. Cooper) ; Ngare Nairobi, W. Kilimanjaro, 4,500 ft. (B. Cooper) ; Shinyanga (N. C. E. Millar). -British Eechuanaland: Ngamiland (G. D. H. Carpenter) ; Ghanzi, Monfalatseka? Ngamiland, off dog (J. Maurice). - India : Quetta, Baluchi- stan (D. Harrison) ; Nedungadu (P. S. Nathan) ; Arbham, Vizagapatam (R. Senior-White) ; Chip~~upalle, Vizaga- patam (R. S. Patuck) ; Kangra Valley, Punjab (Dudgeon) ; Dehra Dun ; Bangalore ; Ehowali, Kumaon, 5,700 ft. (Imms) ; Allahabad ; Bandra (Javakar) . - Ceylon : Madulsima ; Ban- har (R. Senior-White) ; Trincomali. - Assam : Mungpoo, Reang River (R. Senior-White) . - Indo-China : Than-Moi, Tonkin (H. Fruhstorf er) . - China : Macao (F'. Muir) ; Tshusiung, Yunnan, 1,900 m. (Handel-Mazzetti) ; Tsinan, Shantung (E. Hindle) ; Hanchow, off dogs (Rose) ; Kachek, Hainan Id. (L. Gressitt) ; Peiping ; Yen-Ping. - Manchuoko (Manchuria) : Harbin (Jettmar) .
The preapical bristles of the scutellum vary from 3 to 7 in 143 specimens examined (79 ? and 64 8 ), from 23 10- calities, 126 specimens having 5 or 6 bristles. The several numbers are represented as follows: with 3 bristles: I ( 8 );4b.: 3 (I ~ , 2 8 );5b.: 19 (I1 ? , 8 8 ); 6b.: 107 (60
,47 8 ) ; 7b. 13 (7 9 , 6 8 ). There is apparently no sexual difference in this character. The number from any one locality is too small for further analysis. The bristles are evenly divided in I10 specimens, unevenly in 33, the uneven groups observed being I + 2,2 + 3, and 3 + 4. H. longipennis differs from H. equina in the shape of the tergal plates of the abdomen, the difference being more striking in the male than in the female. In the female of H. longipennis the three median, setulose sclerotized plates are very small, ovate or reniform, the second smaller than either the first or the third, the third much smaller than the anterior pair of subapical (lateral) plates. In the male,
the three median plates are large and ribbon-like, much of the same shape as in the male of H. equina; but the hind- most (or third) plate is connected laterally with a pair of small, attenuated lateral subapical plates (recognizable by their bearing longer setae than the median plate proper). No adequate figures of this species have been published. The claspers (or parameres) of the male genitalia are very similar to those of H. equina.




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80 Psyche [ June-Sept.
Historical Note. - There is every reason to believe that Hippobosca longipennis, the dog-fly of the Near and Far East, was well known by the ancient Greeks and Romans, as it is particularly abundant in the countries bordering the eastern Mediterranean. Many are the references to "kuna- muya" (in Greek) or Cynomya in the classic literature and early scientific writings. Thus, in the Iliad, Ares, the god of war, upbraids Athene: "You dog-fly, why do you $ow strife among the Gods? . . ." (Bk. 21, 394). Elsewhere
(Bk. 21, 421) Athene exclaims: "Now watch that dog-fly [meaning Aphroditel leading Ares through the free-for- all. . . "1 I am also inclined to think that the Greek word "kunoraistai" or dog-destroyers, used in the Odyssea (XVII, 300) and later by Aristoteles, covered the ecto- parasites of dogs in general, hippoboscid flies as well as ticks. Oudemans (1926, Tijdschr. v. Entom., LXIX, Suppl., pp. 49-59) claims that both "kunoraistai" and "kunamuya" were used by the Greeks for dog ticks only (Ixodes reduvk Linnaeus). He is evidently unaware of the abundance of Hippobosca longipennis on dogs in the Orient. It seems most improbable that the Greeks would have called a tick a fly, since they had a special word for ticks ("krotones") and must have been well acquainted with both types of para- sites. Moreover, the hippoboscid attracts more readily the attention and is more loathsome to the layman than the tick, owing to its habit of scurrying about in the fur and of flying from one dog to another or even onto people. Hence the use of the word "dog-fly" as a reviling or scurrilous epithet.2
1The exact dating of the collection of epic poems now called the Iliad and credited to Homer is a matter of speculation. Probably they had more or less crystallized into their present form by the eighth century B. c.
2Oudemans also claims that the "musca?' or flies mentioned by Varro, Columella, Plinius and others as causing sores in dogs, were ticks. But in warm countries certain biting flies, such as Stomoxys and Phlebotomus, may cause true sores on the ears of dogs. Oudemans
is apparently also mistaken in criticizing Albertus Magnus' use of the expression "museas bestiarum. quae dicuntur cynodas sive muscse caninse" (De Animalibus Libri XXVI, 1260). Albertus, in my opinion, alluded correctly to the winged hippoboscids which in southern Europe infest horses and cattle, as well as dogs; the two species being so much a^ike that laymen would naturally call them by the same name. In his second volume, Oudemans (1929, pp. 150-151) is quite elated over



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19391 Notes on Hippoboscidae 81
Dr. Gaines Kan-chih Liu has called my attention to ref- erences to dog-flies in the early Chinese medical literature and has kindly translated some of these for my paper. In the "Chi Twq~ Yeh Yu," by Chow Mi (who lived 1232 to l308), one reads: "A colleague of mine, Chen P'o, of Quo Chang, is an old scholar. His grandson, when three years old, was seriously ill with fever for a week, after which "to" (or small-pox) broke out, the whole body turning black and the lips being icy cold. After all remedies had failed, the grandfather went to the temple to pray God for help. There he met a stranger, who, upon learning of the case, told the grandfather how to cure it. The prescription consisted of seven dog-flies, ground into a powder and taken with wine. The medicine was very effective and the child soon was in good health, the black color disappearing." In the later "Pen Tsao Kang Mu" (1578), by Li Shih-chen, one finds: "Dog-flies live on the body of the dog. They can fly, are yellow and fly-like and have a hard skin. They have a sharp beak and suck the blood of the dog. Formerly they were not known to be used in Medicine; but recently they have been recommended by the Chi Tung Yeh Yu for cur- ing small-pox and by the Yi Fang Da Chien for malaria. It seems to me that they must act like the cattle-lice and the chufoo (or sawbugs). For malaria, the flies, after removal of the appendages, are made into pills with dough. They should be taken the morning of the day an attack of fever is due and the cure will be successful if vomiting is provoked. Another method is to make the flies into pills with wax and take the pills with wine. For small-pox and skin troubles, soak the fly in wine and then take both the fly and the wine." Finally the "Chien Wu" (1582), by Li Su, says: "The dog- flies deposit among the hairs of the dog their nits (puparia), which after molting become flies. They always live on the back of the neck, where they bite frequently and where the dog cannot reach them with its mouth or paws." 3.
Hippobosca fulva Austen. - Through the courtesy of his discovery that the Archbishop Eustace of Saloniki, in his Com- mentary of the Iliad, suggests that the Greek word "kunun~uya" (which he proposes to emend to "kunon~~liya") evidently meant the tick. But this statement proves only that the Archbishop was more profi- cient in philology than in natural history.



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82 Psyche [Jane- Sept.
the late Major Austen, I was able to examine the holotype and paratype at the British Museum. This species, which has not been figured, is close to H. longipennIS, the most im- portant differences being given in the key. In addition, the vertex is somewhat narrower than in longipe%ms, with the inner margins slightly converging toward the occiput; the postvertex is shorter; the inner orbits (or parafrontalia) are narrower and of more uniform width throughout; the fronto-clypeus also narrower. The insect is mostly reddish- yellow ; but scutellum, postvertex and fronto-clypeus are al- most wholly pale ivory-yellow. I have seen also a male from Tanganyika Territory (West shore of Lake Manyara) . This sex ia almost exactly like the female. It has eight strong pale-colored preapical bristles on the acutellum (also present in both types), placed in a single row and widely divided into two groups of four each. The structure of the abdomen is not known, but is probably similar to that of equina and Longipennis, with minor differences in the rela- tive size of the median tergal plates.
In the mate of fulva examined, the parameres of the genitalia are similar to those of e* and longipennis, but the terminal point is blunter. H. fidva and H. wia,rtiå´nagZi are the smallest members of the genus.
4. Hippobosca vankgata Megerle von Muhlfeld, 1803, (actual date of publication!), Appendix ad Catal, Insect. Nov. 1802 Viennae Austriae Vendita, p. [14] (unnumbered) (Bengal) .
Synonyms : H. maculata Leach, 1817 ; H. bipartitu Mac- quart, 1843; H. aegyptiaca Macquart, 1843; H. foswlata Macquart, 1843 ;I H. sud-mica Bigot, 1884 ; H. shae Bigot, 1885; H. &psis Bigot, 1885; H. aegtfptiaca var. benga- Lewsis Ormerod, 1895.
H. variegata appears to be the oldest valid name for this species, the date 1823 given by Wiedemann being erroneous (see Schenkling 1935, Arch. Morph. Taxon. Entom., Ber- lin-Dahlem, 11, p. 156). The original description is of the briefest: "ex Beng. Aff. equin. sed rnaj. magisque varieg. (1 Exemplar) ." Yet it is sufficient to validate the name, especially in view of the fact that Wiedemann (1830, 1Macquart's three names should be dated 1843, when the Mem. Soc. Sci. Lille for 1842 were actually published.



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19391 Notes on Hippoboscidae 83
Aussereurop. Zweifl. Ins., 11, p. 603) based the more detailed and fully recognizable description of his Hippobosca varie- gata in part upon Megerlk's specimen.
I have seen, at the University Zoological Museum in Copenhagen, the fly from Tranquebar mentioned by Fabri- cius (1805, Syst. Antliat., p. 338) as a variety of H. equina. It is H. variegata and may well have been the specimen which Wiedemann mentioned from Tranquebar. I have also seen Leach's type of H. maculata at the British Museum. According to the label it came from Bengal. Additional Specimens Examined. - French West Africa : Zinder, Niger River, off cattle and horses (A. Buchanan) . - Gold Coast: Salaga, N. Terr., off cattle (F. J. A. Beringer) ; Yegi, N. Terr.; Accra (J. W. S. Macfie) ; Obuasi, Ashanti (W. M. Graham). -Northern Nigeria: Kaduna (J. J. Simpson) ; Azare (L. Lloyd) ; Zungeru, off horses. - South- ern Nigeria: Lagos (C. B. Philip) ; Olokemeji, Ibadan. - Cameroon : Bamum. - Belgian Congo : Sankuru District, 5' S., 26O E. (A. Yale Massey). - Anglo-Egyptian Sudan: Khor Hanoieit, Port Sudan; Khartoum (S. Hirst) ; Erko- wit, Red Sea Hills, biting man (J. G. Myers) ; near Meshra, Equatoria biting man ( J. G. Myers) . - Egypt : Luxor (Reimoser.) - Ethiopia : Hawash River, W. of Mt. Zaquala, 6,000 ft. (J. 0. Couper) ; Maraquo (0. Kovacs) . - Uganda: Mt. Debasien, 5,000 ft. (A. Loveridge). - Kenya Colony : Nakuru (van Someren) . - Tanganyika Territory : Kigon- sera (J. N. Erth) . - Natal : Durban. - Madagascar : Betsiriry District, west of S. Central Plateau, off mules (F. P. Porter) ; Tsaratanana, N. Central District (W. C. Holden) ; Ampoza (E. I. White) ; Tanovana, Oriental Forest District (between Tamatave and Tananarive. - C. Lamber- ton). - Mesopotamia (Iraq) : Daurah (A. D. Fraser) . - India: Calcutta (Brunetti) ; Bhowali, Kumaon, 5,700 -It. (Irnms) ; Deccan (Fischer) ; Mukteswar, United Prov. (J. D. R. Holmes) ; Pusa, Bihar, off cattle (R. Senior-White) ; Tranquebar. - Ceylon : Banhar (R. Senior-White) ; Matale, off horse (R. Senior-White) ; Luduganga, off cattle (R. Senior-White) ; Habarane, off cattle (R. Senior-White) ; Peradenyia, off cattle (A. Rutherford) ; Dambula, off cattle (L. G. Saunders) ; Hamsantota ; Madulsima ; Diyawa, 4,000 ft. - Assam: Coonoor, off cattle (R. Senior-White) ;



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84 Psyche [ June-Sept.
Baranri (R. Senior-White) . - Timor. - Celebes : Manado, off horse (F. C. Kranevdd).
The preapicd bristles of the scutellum are pale-colored and unevenly developed, usually placed on two irregular rows, the two lateral groups more or less connected in the middle. Counting only the heavy bristles and neglecting the small, hair-like ones, their number varies from 13 to 27 in 95 specimens examined (49 a and 46 a ), from 12 locali- ties, 68 specimens having from 16 to 20 bristles. The speci- mens show the following grouping: with 13 bristles: 2 (all Q );14b.: 1 ( s );15b.:5 (2 s ,3 <t);l6b.:7(6 s ,1 8); 17b.: 15 (7 Q , 8 8 ); 1%: 15 (4 s , 11 8); 19b.: 22(13 Q ,9 s );20b.: 9 (6 s ,3 a);21b.:7 (4 $ ,3 8 ) ;22b.: 2 (1 $ ,1 a);23b.: 3 (2 Q ,1 s ); 24b.: 4 (1 9 , 3 8); 26b.: 2 (all 8 ) ; 27b. : 1 ( 8 ). There is seemingly no sex- ual difference in this character. In one lot of 30 specimens (10 s ,20 8 } from Aden, off cattle, the bristles vary from 13 to 22, but 25 specimens have from 16 to 20 bristles. The bristles are evenly divided between both sides in 37 speci- mens, unevenly in 58, the uneven groupings observed being 6+7,6+8,7+8,8+9,9+10,10+11,11+12,
12 + 13, and 13 + 14.
In H. varkgata, the dorsum of the abdomen bears no median tergal sclerotized plates in either sex. In the female, the anterior pair of subapical (lateral) plates is somewhat larger than the posterior pair. In the male, the anterior pair is very large, so that the two plates nearly touch medi- ally; the posterior pair is very small and hidden in a dorsal view. The abdomen of the female is shown correctly by Ferris (1930, Philippine 31. Sci., XLIII, p. 545, fig. 5). There is no good figure of the male.
The claspers (or parameres) of the male genitalia are straight and rod-like, rather abruptly narrowed about mid- length and then very slender to the pointed tip. 5. Bippobosm rufipes v. Olfers.
Additional Specimens Examined. - Natal : Durban, off
cattle (W. C. C. Pakes). - Cape Province : Van Rhyn's Pass (T.D.A. Cockerell) ; Mihertown near Cape Town (R. E. Turner) ; Waku (J. Brace-Bays) ; Matjeafontein (TS.. E. Turner) ; Erraha (E. Gough). - South West Africa : Tsau,
Great Namaqualand (Poch) ; Otyivarongo, Damamland (de



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19391 Notes on Hippoboscidae 85
Schauensee) ; Windhoek to Gobabis (de Schauensee) ; Okahandja
(R, E. Turner ; J. Ogilvie) ; Otavifontein (K. Jordan) ; Usakos (J. Ogilvie) ; Hoffnung (K. Jordan). - Orange Free State:
North Bank Halt, Norvals Poort (J.
Ogilvie) ; Cotzies' Farm (W. L. Distant). - Bechuanaland Protectorate: Mongalatsela, Ghanzi, Ngamiland, off horse and off steinbok (J. Maurice). - Southern Rhodesia: Vic- toria Falls (R. Lowe Thompson). -Northern Rhodesia: Lunda, near Congo border (H. S. Evans). - Portuguese West Africa: Benguela, off cattle (W. C. C. Pakes). - Tan- ganyika Territory: Mt. Meru, 4,500 to 5,000 ft., off eland (I3 Cooper) ; Ngare Nairobi, W. Kilimanjaro, 5,000 ft. (B. Cooper) ; Ngaserai, W. Kilimanjaro, 3,000 ft. (B. Cooper). The number of heavy, black preapical bristles of the scutellum is quite variable in this species and apparently shows no sexual difference. In 95 specimens examined (69 Q and 26 8 ), from 14 localities, the total number va- ries from 12 to 23, but 82 specimens have from 14 to 20 bristles. The specimens are distributed as follows: with 12 bristles: 2 (I Q , I 8 ); 13b.: 5 (4 Q , I 8 ); 14b.: 10 (8 Q ,2 8); I5b.: I0 (all Q ); 16b.:7 (6 Q ,I 8 ); 17b.: 17 (13 Q ,4 8 );18b.:18 (I1 Q , 7 8 ); 19b.: 12 Most localities are represented by one or a few specimens. In a series of 67 specimens taken off cattle at Windsorton, Cape Province, the proportion is as follows: with 13b.: 3 (all 9 ) ; 14b.: 4 (3 Q , 1 8 ) ; 15b.: 9 (all Q ) ; 16b.: 5 (all
Q ); 17b.: 12 (9 ? , 3 8 );18b.: 15 (I0 Q , 5 8 ) ; 19b.: 8 (5 2 ,3 8 );20b.:5 (2 Q , 3 8 );21b.: 5 (4 9 ,I 8 ); 22b. : 1 (
8 ) . The bristles are more often unevenly divided between both sides (unevenly in 58 specimens, evenly in 371, the two groups only narrowly divided, so that the row is fairly continuous. The uneven groupings observed were 6 +7,7+8,8 +9,6+8, 7+9,8+10,9+10,9+11,
10 + 11,9 + 12, and I1 + 12.
The structure of the abdomen of H. rufipes is similar to that of H. variegata. It is correctly shown for the female by Austen (1909, Illustr. African Blood-Suck. Flies, Pl. XIII, fig. 100).
The parameres of the male genitalia are straight and



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86 Psyche [ June-Sept.
rod-like, abruptly narrowed at basal third, beyond which they are thicker than in H. variegata and blunter at the pointed apex.
6.
Hippobosca hirsuta Austen. -1 have seen the types of the typical form and the var. neavei Austen at the British Museum.
Additional Specimens Examined. - Uganda : near Lake Albert (H. Hargreaves) . - Nyasaland : Akamanga, Run- yinya River, N. Nyas3, off waterbuck (J. B. Davey). The preapical bristles of the scutellum are much weaker in this species than in most other members of the genus, and pale-colored. In the few specimens seen their total number varies from 14 to 18 (I with 14, I with 15, I with 16, and 1 with 18, for the males; I with 14 and 1 with 16, for the females) .
The structure of the abdomen is similar to that of H. variegata in both sexes.
The parameres of the male genitalia differ from those of all other species of the genus. They are broad and thick, beam-like, slightly wider basally and apically in side-view. The apex is straightly truncate, shall~wly and evenly emar- ginate, so as to produce two blunt edges, one dorsal and one ventral.
7. Hippobosca martinaglia Bedford, 1936, Onderste- poort Jl. Vet. Sci. An. Ind., VII, pt. I, p. 67, fig. (on p. 68) ( 9 8 ; off Impalla, Aepyce~os melampus (Lichtenstein) ; Bar R. Ranch, Swaziland, South Africa).
In the absence of specimens, I have inserted this species in my key on the assumption that there are set2 on the basal section of the third longitudinal vein and that only one of the pulvilli is well-developed, neither character being mentioned by the author. From the figure and description, it appears to be closest to H. hirsuta, likewise found on antelopes. It differs in the smaller size, the few short black setae of the mesonotum, the few preapical bristles of the scutellum, the wider frons, the narrower inner orbits or parafrontalia (less than half the width of the mediovertex or frontalia), and the short, semi-elliptical postvertex. The second longi- tudinal vein is described as "long", but the figure shows it to be "short", reaching the costa a short distance beyond the



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tip of the first longitudinal vein, so that the last ~ection of the costa is about six times the length of the penultimate section. No median tergal plates are shorn in the figure of the female. The co1oratjon seems to fit some of my speci- mens of H. hngipmnis cape mi^)^ but the second longid tudinal vein is much shorter than in that species. The venation also separates H. raahinag1.h from H, fdvu, a fly of about the same size, likewise found on antelopes. !l%e original description is reproduced) as it appeared in a periodical inacces~ible to most entomologists: "A small species ; length of wing 4.5 mrn. Head about as wide at the occiput as at the fronto-dypeus, reddish-brown) the frontal stripe slightly darker; posterior margin of head fringed with minute) thick-set sets, and a long seta at the base of each eye; palpi dark brown) clothed with short setae of the same colour. Thorax reddish-brown, 6th a median dark band extending backwards ahmost to the transverse suture ; this bard is forked poshriorly, usually moxe so than in the figure, and in one specimen is completely divided down the middle by a narrow line; on eaeh side of the posterior por- tion of the median hand there is a dark transverse band, On each side beneath the transverse suture there is a nax- row transverse dark band, and beneath this 8 8maH tri- angular spot, which is usually indistinct and may be absent. At each latero-anterior angle there are two short set=, one on each side slightly distad and nearer the meson ; on each side above the base of the wing there are three very short, thick-set black sek, two moxe similar set= slightly above them and near the meson) and two larger set= slightly above them and near the meson, and two larger set= below them ; on eaeh side on the posterior margin there are five sets?. Scutelhm yellowish-white, fringed with short and a few long sek. On the venter there is a vertical dark band on each side between the fore and mid cox=. Legs pale red- dish-brown, sparsely clothed with set=; those on the t h j ~ and tarsi darker. Ungues black. Abdomen reddish-brown with numerous pale s e k Wings hyaline with pale reddish- brown veins and short dark sek on the costa, Second longi- tudinal vein (R2 + 3) long, reaching beyond the apex of the first longitudind vein (RI ) , but not extending to the ante- rior cross-vein. This new species can be easily recognised



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88 Psyche [ June-Sept.
by its pale colour and dark markings on the thorax and venter between the fore and mid coxae; also by the short thick-set sek on the thorax and pale scutellum." 8. Hippobosca struthionis Janson. -1 have seen the type from Mt. Stewart, Cape Province, at the British Museum.
Additional Specimens Examined. - Kenya Colony : Simba, 3,350 ft. ; Makumbu ; Ukamba; Machakos; Athi Plains, off horse; Merifano (McArthur) ; Taveta, off dog (C. W. Wood- house). - Tanganyika Territory : Tabora? off ostrich (J. Rodhain) ; north of Tarengere River (W, A. Lamborn) ; Sanga River, Muruangani. - Transvaal : Pretoria ; Deel- fontein; near Limpopo River, N. Transvaal, off ostrich (R. A. Cooley) . - Cape Province : Philipsdale, Worcester ; Hopetown; Campbell ; Cradoek (Miss J. Brincker) ; Erreha (E. Gough) . - Beehuanaland : Ghanzi, Ngamiland (J. Maurice). - South West Africa : Otavi (J. Ogilvie) ; Aus (R. E. Turner) ; Greater Spitzkopje near Usakos, Damara- land (de Schauensee) .
This species is unusual in that the preapical bristles of the scutellum are crowded together in two widely separated groups, placed at the extreme sides; also in that the bristles are more numerous in the male than in the female. In 23 specimens (13 Q and 10 8 ) examined, from 4 localities, the bristles varied in the females from 4 to 12, 9 of the 13 Q having from 5 to 7; in the males, from 8 to 15, 7 of the I0 8 having from 9 to 12.
The grouping was as follows : with 4
bristles: 2 ( Q ); 5b.: 3( Q ) ; 6b.: 3 ( Q ); 7b.: 3 ( Q ); 8b.: I ( 8 );9b.:4 (I Q ,3 8 );lib.: 2 ( 8 );IZb.: 3 (I 9 ,2 8 );18b.: I ( 8 ); 14b.: I ( 8 ) ;15b.: I ( 8 ). The bristles were divided into even groups in 8 specimens, unevenly in 15? the uneven groupings observed being 2 +- 3, 3 + 4, 4 +
5 + 6, 5 + 7, 6 + 7,and7 + 8.
The structure of the abdomen of H. struthionis is similar to that of H. variegata in both sexes. The species has not been figured adequately.
The parameres of the male genitalia are slender, rod-like, gradually narrowed from the base to the pointed, sharp apex, and very similar to those of H. equina. 9.
Hippobosca camelina Leach. -1 have seen Leach's type, at the British M'useum.




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19391 Notes on Hippo boscidue 89
Additional Specimens Examined. - Morocco : Debdou- Taourirt (E bner ) ; Tendrara (Ebner) . - Mauretania : Be- tween Kiff a and Tidjidja, off camels (Mrs. Mary Steele) . - Algeria : Biskra (Rothschild and Hartert) ; Ain Sefra (Rothschild land Hartert) ; Touggourt (Rothschild and Hartert) ; Oued Nqa, Mzab country (Hartert) ; Zahrez Gharbi (Zerny) . - Tunis : Gabes (Mik) ; Tozeur (G. F. de Witte) . - Libya :
Tripoli (Klaptocz) ; Dernah (Klaptocz) ; Bengasi (Klaptocz) . - Egypt : Old Cairo, off horse (Effla- toun Bey) ; Gebel Elba, S. E. Desert (Tewfik) ; near Pyra- mids (Reimoser) . - Palestine : Jericho (P. A. Buxton) ; Jerusalem (P. Barraud). - Arabia : Djedda (13. W. Whyte) ; Tuwaiq, Riyadh (H. St. J. B. Philby) ; Keshin (Hein). -British Somaliland: without more definite 10- cality (C. L. Collenette). -Kenya Colony: Lodwar, Tur- kana; Marsahit, Rendili Nyoro, N. Frontier District (C. A. Neave) ; near mouth of Kallilokwelli River, Lake Rudolf (E. B. Worthington) . - Anglo-Egyptian Sudan : Khor Hanoieit, Port Sudan; Erkowit, Red Sea Hills, biting man (J. G. Myers).
H. cumelinu is in many respects the most aberrant mem- ber of the genus, particularly in the arrangement of the scutellar bristles and in the structure of the abdomen, both features showing remarkable sexual diff erences. If few specimens were examined, one might be led to think that two species are confused under the name camelinu. For- tunately I was able to study a series of 20 specimens (I0 Q and 10 8 ), collected in one locality of British Somaliland, removing all doubt in the matter.
In the male the scutellar bristles are preapical, placed as in the other species of the genus. In 14 males examined, from 4 localities, their number varies from I1 to 22, with the following groupings:
I1 bristles : I ; 12b. : 4 ; 14b. : 2 ;
15b. : 4 ; 16b. : I ; 17b. : I ; 22b. : I. They are divided into two even groups in 6 males, unevenly in 8. In the female the bristles are discal, placed near the middle of the scutellum and much more spaced. In 17 females, from 6 localities, their number varies from 3 to 8, as follows: 3 bristles: I ;
4b. : 6 ; 5b. : 4; 6b.: 3 ; 7b. : 2 ; 8b. : I. They are evenly di- vided in 10 females, unevenly in 7.
In the female the dorsum of the abdomen is entirely soft



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90 Psyche [ June-Sept.
and extensible behind the usual basal tergal sclerite; post- eriorly one finds on the extreme sides two pairs of subapical strongly sclerotized plates, bearing long bristles? the ante- rior pair being much larger than the posterior pair. In the male there is immediately behind the basal tergal sclerite a large ribbon-like median plate? somewhat triangular medial- ly; much farther back? also a pair of very small median plates, rather far apart; two pairs of subapical plates are present? but the posterior pair are very small and hidden from view in a dorsal aspect. The abdomen is fairly eor- rectly drawn for both sexes by Massonnat (1909, Ann. Univ. Lyon, N.S.? CXXVII19 Pl. III? figs. 24 and 25). The parameres of the male genitalia are peculiar. They are rod-like seen from above? but in side view more or less boat-shaped; the basal two-thirds are wide? with an evenly convex lower margin; the slender apical third is curved up- ward and ends in a blunt, somewhat knob-like point. The outer surface is raised into a median? curved? blunt ridge? running the whole length.




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