J. Bequaert.
Notes on Hippoboscidæ. 13. A Second Eevision of the Hippoboscinæ.
Psyche 46:70-90, 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.