J. Bequaert.
Notes on Hippoboscidae.
Psyche 40:68-82, 1933.

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Psyche
[June
NOTES ON HIPPOBOSCIDAE
4. ON THE LARGER SPECIES OF LYNCHIA WEYENBERGH (OLFERSIA OF AUTHORS; ICOSTA SPEISER;
ORNITHOPONUS ALDRICH)
Department of Tropical Medicine,
Harvard Medical School, Boston, Mass.
On size alone, most of the species of Lynchia may be di- vided rather readily into two groups.
Although this ar-
rangement disregards true relationship, I shall use it for the purpose of this paper,' which discusses only the larger forms of the genus. These are 10 mm. or more in length, as measured from the tip of the fronto-clypeus to the tips of the folded wings. The length of the wing, a much more reliable standard, varies from 6 to 8.5 mm. in the specimens I have seen. L. pilosa (Macquart) is the smallest member of the group known to me. L. penelope Weyenbergh, the un- recognized genotype, also belongs to this group. I have seen specimens of the six species included in the subjoined key. Having ,studied much material, I am in- dined to regard most of the other names proposed for large Lynch&, as synonyms and I have attempted to treat them accordingly. In many cases the identity of these names must remain open to question, until the original specimens can be examined. I have seen the types of Hippobosca bu- bonk Packard and Hippobosca nigra Perty only. 1. Mesonotum and scutellum covered fairly uniformly with soft, short hairs, directed backward; scutellum with- out median groove. Frons very wide, with the sides distinctly converging below; postvertex very short and wide, about one-fourth of the length of the medio-ver- tex; inner orbital bristles of frons very numerous, in several irregular rows;
palpi short, slightly shorter
Pu&e W:68-82 (1933). hup Ytpsychu einclub orgHW&OM html



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Notes on Hippoboscidae 69
than the fronto-clypeus. Basal two-thirds of wing almost destitute of setulae on the upper side; second basal cell (M) only about one-third the length of the first (R) ; subcosta (SC) complete, ending in costa. Wing 6 to 6.5 mm. long. African species ........................ ....................................................... L. pilosa ( Macquart) .
Mesonotum almost bare, except for patches of hairs be- hind the humeri and before the scutellum; scutellum with a distinct median depression, groove or furrow, the disk bare except for a pair of lateral setae and an apical fringe of soft hairs.
Palpi as long as or longer
than fronto-clypeus. Upper side of wing mostly covered with setulse, except the axillary cell (2d An) or this cell and the posterior fourth to half of the anal ........................................................ cell (Cu + 1st An)
2.
2. Upper side of wing mostly covered with setulse; only the axillary cell (2d An) bare. Inner orbital bristles of frons fairly numerous, mostly in one row, but a few forming an incomplete second row. Subcosta (SC) complete, ending in costa. Frons nearly twice as wide as an eye, with subparallel sides. Postvertex rather short, the anterior margin longer than each of the ............................................ sides. Wing 7 to 8 mm. long
................ L. palustris (Lutz, Neiva and Costa Lima) . Axillary cell and posterior fourth to half of anal cell bare ................................................................................ 3. 3. Bristles on inner margin of orbital plates few in num- ber, placed mostly in one, irregular row. Subcosta (or
auxiliary vein ; SC) incomplete, not ending in costa.. 4. Inner orbital bristles of frons numerous, placed in more than one row. Subcosta (SC) complete, ending in costa. Frons about one and a half times to twice as wide as an eye; inner orbits distinctly diverging to- . ward vertex .................................................................... 5. 4. Inner orbits subparallel. Smooth postvertex rather long, semi-elliptical, with broadly rounded anterior angles, the anterior margin about as long as the sides; with a rudimentary ocellus in a median pit. Wing 6.5 to 7 mm. long ....................................... L. fusca (Macquart) .



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70 Psyche [June
Inner orbits very distinctly diverging toward the vertex. Smooth postvertex rather short, transversely elongate, the anterior angles abruptly rounded off, the anterior margin rather straight and longer than each of the sides; without median pit and rudimentary ocellus. .
Wing 7 to 8.5 mm. long.. .......... ..L. americana (Leach) 5. American and Pacific species. Postvertex rather long ; the anterior angles broadly rounded off, the anterior margin about as long as each of the sides. Wing 7.5 to 8.5 mm. long ....................................... .L. nigra (Perty) . African species. Postvertex rather short, more trans- verse; the anterior angles more abruptly rounded off, the anterior margin much longer than each of the sides. Wing 8 to 8.5 mm. long ...... ..L. dukei (Austen). In the foregoing key the terms "postvertex" and 4'fro~nto-clypeus'y are used, as defined by Jobling. The postvertex corresponds to the "vertical plate" or "vertex" of most authors, being the hard, smooth area in the upper part of the frons, limited behind by the occipital margin. Lynchia pilosa (Macquart)
Olfersia, pilosa Macquart, 1843, M6m. Soc. Sci. Lille, (1842), p. 434- (no sex ; Africa, without more definite lo- cality; but since the type was collected by Delalande, it must have come from South Africa) ; 1843, Dipt. Exot., 11, pt. 3, p. 277. Neave, 1912, Bull. Ent. Res., 111, pp. 317, 320 and 322. Morstatt, 1913, Der Pflanzer, IX, p. 509. Lynchia pilosa Falcoz, 1930, Encyclop. Entom., Diptera, V, p. 48. Bedford, 1932, 18th Rept. Director Vet. Serv. Animal Ind. Union S. Africa, p. 424.
SPECIMENS EXAMINED.-~~~~~~ Free State : Hoopstad, one female (H. Brauns) .-Abyssinia : Ouache (= Hawash River ?) , one male (Paris Museum).
This remarkable species is readily recognizable by the peculiar pilosity of the thorax and the hyaline, extensively bare wings.
The present location of the type is unknown. The hosts are species of bustards, according to G. A. H. Bedford (1932), who records L. pdosa off the Kori, Choriotis kori



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19331 Notes on Hippoboscidae 71
(Burchell) , in Zululand ; the White-quilled Black Knor- haan, A frotis atra af raoides (A. Smith), in Transvaal ; and the Greater Bustard, either Choriotis kori struthiunculus (Neumann) or Neotis cafra jacksoni Bannerman, in Kenya Colony.
Lynchia palustris
(Lutz, Neiva and da Costa Lima)
Olfersia palustris Lutz, Neiva and da Costa Lima, 1915, Mem. Inst. Osw. Cruz, VII, p. 183, PI. XXVIII, fig. 4 (no sex; off Herodias egretta, Tigrisoma brasiliense, Can- croma cochlearia and Harpiprion cayennensis, State of Piauhy ; off a white heron, Rio de Siio Francisco, Brazil : and off Ardea socoi, Lassance, State of Minas Geraes). Ad. Lutz, 1928, Est. Zool. Paras. Venezolanas, p. 9. ? Olfersia americana Massonnat, 1909, Ann. Univ. Lyon, N. S., CXXVIII, p. 304, PI. V, figs. 40-42 ( Q ). Not of Leach.
? Ornithoponus massonnati Falcoz, 1926, Faune de France, XIV, Dipteres Pupipares, p. 31, figs. 28-29 ( Q ; off Plata- lea leucorodia; region of the Dombes, Ain, France). SPECIMEN ExAMINED.-B~~~~~~ Congo: Mongende, one male ; off a Cormorant, Phalacrocorax africanus (Gmelin) , April 14, 1921 (H. Schouteden).
I am unable to separate this African specimen from L. palustris, commonly found in South America on wading birds. The species appears to be most closely related to L. hirsuta Ferris, of North America; but that fly is much smaller (wing 5 mm. long), has the basal cell relatively shorter, three long setae on each half of prescutum, and a long prescutellar on each side. I find no such setae in my specimen of L. palustris. L. botaurinorum Swenk, which I have seen from Brazil, as well as from North America, likewise differs from L. palustris in the smaller size (wing 5 mm. long).
L. palustris is readily differentiated from L. ardege (Mac- quart) and related species, found more commonly on wad- ing birds, by the larger size, the much wider fronto-clypeus, and the absence of setae over an extensive median area on the dorsal face of the abdomen.




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72 Psyche [June
SYNONYMY.-The name Ornithoponus massonnati was introduced by Falcoz for the fly from France which Mas- sonnat (1909) described and figured, as "Olf ersia ameri- cana," (the type probably in the collections of the Uni- versity at Lyons). Ferris and Cole (1932) were quite jus- tified in concluding that Massonnat's insect was probably "an entirely distinct species" from "americana." Among the larger species of Lynchia (Massonnat gives the length of the wing as 8.2 mm.) known to me, Massonnat's description agrees best with L. palustris, although the hairs of the in- ner orbital plates and the peculiar setulose covering of the wing membrane are not mentioned by him nor by Falcoz. There are some features of Massonnat's drawings, such as the very wide frons, that do not fit L. americana, though some of them might be due to inaccuracies (there are strik- ing differences between the drawings of the head in figures 40 and 42, particularly in the outline of the postvertex and the trend of the sides of the frons). In addition, Masson- nat's fly was taken off a white spoonbill, Plcuhalea leucorodia, and thus far I have never seen specimens of true L. ameri- cana taken from a wading bird. In view of the fact that L. palmtris has now been found on a cormorant in Africa, it seems highly probable'that Ornithoponus massonnati is the same species.
Lynchia americana (Leach)
Feronia, americana Leach, 1817, Gen. Spec. Eprobosc. Ins., p. 11, PI. XXVII, figs. 1-3
(no sex; no host; Georgia,
North America). Austen, 1903, Ann. Mag. Nat. Hist., (7) XII, p. 264 (type at Brit. Mus.). Lutz, Neiva and da Costa Lima, 1915, Mem. Inst. Osw. Cruz, VII, p. 188. Olf ersia americana Wiedemann, 1830, Aussereurop. Zweifl. Ins., 11, p. 606. Macquart, 1835, Hist. Nat. Ins. Dipt., 11, p. 641. T. W. Harris, 1835, in Hitchcock, Rept. Geol. Miner. Zool. Bot. Massachusetts, 2d Ed., p. 600. Walker, 1849, List Dipt. Brit. Mus., IV, p. 1141. Osten Sacken, 1858, Cat. Dipt. North America, p. 86. v.d. Wulp, 1867, Tijdschr. v. Entom., X, p. 128. Osten Sacken, 1878, Cat. Dipt. North America, 2d Ed., p. 213. C. W. Johnson, 1900, 27th Rept. New Jersey Bd. Agric., (1899), Suppl.,



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19331 Notes on Hippoboscidae 73
p. 699. Aldrich, 1905, Cat. North Amer. Dipt., p. 655. Speiser, 1907, Ent. News, XVIII, p. 104. Swenk, 1916, Jl. New York Ent. Soc., XXIV, p. 130. W. E. Britton, 1920, Conn. Geol. Nat. Hist. Surv., Bull. 31, p. 210. Cole and Lovett, 1921, Proc. California Ac. Sci., (4) XI, p. 344. W. T. Davis, 1922, Proc. Staten Island Inst. Arts Sci., I, p. 65. C. W. Johnson, 1922, Psyche, XXIX, p. 83. Ormthoponus americanus Aldrich, 1923, Insecutor Insc. Menstr., XI, pp. 77 and 78. C. W. Johnson, 1925, Proc. Boston Soc. Nat. Hist., XXXVIII, p. 99; 1925, Bull. Northeast. Bird-Banding Assoc., I, p. 52; 1925, Occas. Pap. Boston Soc. Nat. Hist., VII, p. 293. Gross, 1925, The Auk, XLII, p. 431. C. W. Johnson, 1927, Bull. Bos- ton Soc. Nat. Hist., No. 42, p. 15. 0. A. Johannsen, 1928, in Leonard, List Insects New York, (1926), p. 868. Lynchia americana Falcoz, 1930, Encyclop. Entom., Dip- tera, V, p. 48 (in part?). C. W. Johnson, 1929, Bull. Northeast. Bird-Banding Assoc.', V, p. 52. Ferris, 1930, Canad. Entom., LXXII, p. 67, fig. 5A, D, and F. Hippobosca tubonis Packard, 1869, Guide to the Study of Insects, 1st Ed., p. 417 (no sex; off Great Horned Owl; Massachusetts). Copied without change in all later edi- tions.
? Ornithomyia villaclse Dugte, 1887, La Naturaleza, Mexico, (2) I, pt. 1, p. 20, PI. 111, fig. 3 (no sex; off Buteo calurus and B. bairdic somewhere in Mexico; no definite locality given).
POlfersia villudse van der Wulp, 1903, Biol. Centr. Amer., Diptera, 11, p. 430. Aldrich, 1905, Cat. North Amer. Dipt., p. 656.
The following references to "Olfersia americana" are doubtful or erroneous.
Olfersia americana Ferris and Cole, 1922, Parasitology, XIV, p. 194, figs. 11-12 ( ? ; off Tyto alba pratincola; San Bernardino, California). To judge from fig. 11, the frons is distinctly less than twice the width of an eye and has sub- parallel sides; while the postvertex is rather long in the middle, with broadly rounded angles. Most probably this specimen was what I call L. fusca (Macquart).



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74 Psyche [June
LynchuS, amenCaw Ferris, 1927, Canad. Entorn., LIX, p. 248, figs. 2 and 3 (fig. 3 erroneously labelled "L. hirsuta"; corrected by Ferris, 1929, Ibidem, LXI, p. 285) (9 ; off
Oh asio; Monticito, California). The same remark ap- plies to this as to the foregoing record. 0-thoponm umericanus C. W. Johnson, 1924, Zoologica, New York, V, No. 8, p. 91 (off Buteo galapagoensis; Sey- mour Bay, Indefatigable, Galapagos).
I have seen this fly,
which is L. mgra (Perty) . Curran's 0. amehcams, from Santa Cruz, Indefatigable, (1932, Nyt Mag. Naturvid., LXXI, p. 3661, probably also was L. nigra, but I have not seen the specimen.
Olfersia americana C. W. Johnson, 1896, Proc. Ac. Nat. Sci. Phila., p. 340 (off Screech Owl, Otus asio floridanus; St. Petersburg, Florida). I have seen this specimen, which is L. fusca (Macquart ) .
Olfwsia, amerZaanu C. W. Johnson, 1918, Bull. Amer. Mus. Nat. Hist., XXXII, p. 90 (St. Augustine and Miami, Florida). These specimens, seen by me, are L. fwca (Macquart) .
SPECIMENS EXAMINED.-L. mericana is perhaps the moat common of the North American hippoboscids. I have seen over one hundredspecimens from the following localities. Ontario : Point Pelee.-Nova Scotia : Annapolis.-Maine : Kittery Point.-New Hampshire: Candia ; Hamptm-Ver- mont : Dunmore Lake.-Massachusetts : Boston ; Wenham ; Middleboro ; West Tisbury ; Framingham ; Monterey ; Do- ver ; Sherborn ; Princeton ; Chicopee.-Connecticut : Liberty Hill,-New York : Long Island ; Staten Island ; Freeport ; Yankee Lake, Wurtsboro, Sullivan Co. ; Austerlitz ; Ithaca ; West Point; Albany; Sparta; Clarksville; Albany Co. ; Baldwinsville ; Camel.-New Jersey : Ramsey ; Haddan- field ; Stag Lake, Sussex Co. ; Kittatinny Mts.-Pennsyi- vania: Eschol ; Penfield ; Dauphin Co. ; Philadelphia ; Shir- leysburg ; Pike Co. ; Harrisburg.-Virginia : Urbana.- North Carolina : Raleigh.-South Carolina ; Charleston ; Dewees Island; Society Hill,-Florida: St. Augustine.- Ohio : Licking- Co. ; Crestline; Wauseon.-Indiana : Indian- apolis.-Illinois : Warsaw.-Wisconsin : W. Spring* Green ; Ripon.-Iowa : Iowa City.-Kansas : Clay Co. ; Douglas Go.



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1.9331 Notes on Hippoboscidae 75
-Texas : Dallas.-Nevada : without more definite locality. -Mexico : Grito (in which State?) .-In addition there' are reliable published records from Washington, D. C., Georgia, Nebraska and Kentucky.
The host list is a very large one, including owls, hawks and other birds of prey, as well as grouse: Great Horned Owl, Bubo virginianus (Gmelin) , (very common on several subspecies) ; Barred Owl, Strix varia Barton ; Long-eared Owl, Asio zuilsonianus (Lesson) ; Barn Owl, Tyto alba pratincoh (Bonaparte) ; Screech Owl, Otus asio (Lin- nseus) ; Pigeon Hawk, Falco columbarius Linnaeus; Sharp- shinned Hawk, Accipiter velox (Wilson) ; Red-tailed Hawk, Buteo borealis ( Gmelin) ; Broad-winged Hawk, Buteo platypterus (Vieillot) ; Rough-legged Hawk, Buteo lagopus sancti-jo~hannis (Gmelin) ; Red-shouldered Hawk, Buteo lineatus (Gmelin) ; Western red-tailed Hawk, Buteo bore- alis calurzis (Cassin) ; Marsh Hawk, Circus hudsonius (Linnaeus) ; Cooper's Hawk, Accipiter cooperi (Bona- parte) ; Golden Eagle, Aquila chrysaetos (Linnaeus) ; Gos- hawk, Astur atricapillus (Wilson) ; and Ruffed Grouse, Bonasa urnbellus (Linnaeus). At the Mus. Comp. Zool. there is also an old specimen, without locality, labelled "off wild turkey." Sometimes the parasite is very abundant. C. W. Johnson records a Great Horned Owl that yielded 32 flies and some pupae, the latter hidden in the ears. Owing to frequent confusion with related species (par- ticularly with L. fzisca), some previously published records of hosts and localities are unreliable. Two old specimens labelled "Nevada" at the Mus. Comp. Zool., and one speci- men from St. Augustine, Florida, in the C. W. Johnson Col- lection, are true L. americana. But all other specimens I have seen from west of the Rockies and from Florida were L. fusca.
Published records from Oregon (Cole and Lovett, 1921, Proc. California Ac. Sci., (4) XI, p. 344; Falcoz, 1930, Encyclop. Entom., Diptera, V, p. 48), and California (C. W. Johnson, 1922, Psyche, XXIX, p. 83; in addition to those mentioned before), are all open to question. There is one definite record of L. americana from Mexico, but how widely it is distributed there is unknown. Falcoz' indica- tion "Columbia" refers to Washington, D. C., not to British



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76 Psyche 1 June
Columbia nor to Colombia. There is as yet no conclusive evidence that L. americana occurs in Europe, as shown in the discussion of L. palustris.
' SYNONYMY.-The type of Feronia americana (Leach) is at present at the British Museum and it is reasonably cer- tain that it agrees with the characters here given for the species.
That Hippobosca bubonis (Packard) is identical with L. americana
(Leach), was recognized by Osten Sacken as early as 1878. The type, which I have seen at the Museum of Comparative Zoology, shows the correctness of this syn- onymy.
From the description alone it is difficult to recognize Ornithomyia villa& Dug&, (the type of which is probably lost), although there can be no doubt about its being a Lynchia. For a time I believed that it might be L. nigra (Perty), and I have named specimens of L. nigra, in some collections, "villa&." Recently, however, I received, from the Department of Entomology of Kansas University, a fly taken off Buteo borealis calurus at Grito, Mexico, and bear- ing a manuscript label "resemble & Ornithornyia villa& A. Dug." This specimen agrees in every respect with the North American L. americana. I am, therefore, listing villadse, provisionally at any rate, in the synonymy of americana, pending the examination of further Mexican material.
Lynchia fusca (Macquart)
Olfersia fusca Macquart, 1845, M6m. Soc. Sci. Lille, (1844), p. 346 (no sex; Nouvelle Grenade = Colombia) ; 1846, Dipt. Exot., Suppl. I, p. 218. Speiser, 1902, Zeitschr. Syst. Hym. Dipt., 11, p. 177 (description of type). Lutz, Neiva and da Costa Lima, 1915, Mem. Inst. Osw. Cruz, VII, pp. 182, 189 and 194 (doubtful records). Bau, 1929, Zoolog. Anzeiger, LXXXV, p. 10.
Lynchia fzisca Falcoz, 1930, Encyclop. Entom., Diptera, V, p. 47.
? Olfersia macquartii Rondani, 1878, Ann. Mus. Civ. Ge- nova, XII, p. 160 (no sex; New Grenada = Colombia; based upon a specimen received as "fusca" from Mac-



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19331 Notes on Hippoboscidae 77
quart, but claimed to be of a different species). Lutz,
Neiva and da Costa Lima, 1915, Mem. Inst. Osw. Cruz, VII, p. 190.
? Olfersia wolcotti Swenk, 1916, Jl. New York Ent. Soc., XXIV, p. 132 (no sex; off Buteo platypterus; Ann Arbor, Michigan).
SPECIMENS ExAMINED.-F~o~~~~: St. Augustine, off a Screech Owl, Otus asio floridanus (Ridgway) (C. W. John- son) ; Miami (P. Laurent) .-Colorado : one female without more definite locality (Ks. Univ. Ent. Mus.) .-California: several specimens off Burrowing Owl, Speotyto cunicularia hypogea Bonaparte, and off Californian Screech Owl, Otus asio mccalli (Cassin), without more definite locality (H. Edwards.-M. C. Z.) ; San Jose, Santa Clara Co., off Great Horned Owl, Bubo virginianus pacificus Cassin, and off Barn Owl, Tyto alba tuidara J. E. Gray, three specimens (G. A. Allen) .-Also several specimens, without localities, in the old Osten Sacken Collection at the M. C. Z. The specimens from Florida, here listed as L. fusca, were recorded by C. W. Johnson (in 1895, 1913, and 1922) as L. americana. As suggested above, most previous records of L. americana from California and Oregon were probably based on L. fusca.
The differences between L. fusca and L. americana, men- tioned in my key, may seem trifling. Yet they appear to be constant and in the large series of both species studied I have seen no specimen of doubtful identity. A repeated and careful study of my material has failed to disclose other reliable characters. The following peculiarities may, how- ever, be of some value. (1) The frons is usually narrower in L. fusca than in L. americana, although the width seems to be variable in both species; in most fusca it is less than one and a half times as wide as the eye; in most americana it is one and a half times to twice as wide as the eye. (2) The apical, flaring arms of the fronto-clypeus appear to be less spreading in fusca than in americana. (3) The color of body and legs is as a rule more uniform and darker mahog- any brown in fusca, lighter and blotched in americana, while the wings of fzisca lack the slight yellowish tinge of amenkana.




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78 Psyche [ June
SYNONYMY.-T~~~ the flies here called L. fusca are spe- cifically distinct from L. americana can hardly be doubted. A specimen without locality, received from the Zoological Institute of Halle a.S. (through Dr. Vult Ziehen), had been named ('fusca Macquart" by the late A. Bau and this identi- fication appears to be correct. Macquart's species could not possibly be recognized from the original description. For- tunately, Speiser has fully redescribed the type (from the Bligot Collection, now in the possession of Mr. J. E. Collin), and his account tallies well with the specimens listed above. I am inclined to regard Olfersia wolcotti (type in the Dept. of Zoology, University of Nebraska at Lincoln, Ne- braska), as a synonym of L. fusca. Swenk's brief descrip- tion agrees with my specimens, except for his statement that the frons is "not wider than the breadth of an eye." In all my specimens of L. fusca, the frons is distinctly, though often only very slightly, wider than an eye. Falcoz (1930) lists as synonyms of L. fusca both Olfersia macquartii Rondani and (with doubt) 0. angustifrons v.d.