Shouldered Hawk and Aplomado Falcon from Quaternary Asphalt

MARCH 2003 SHORT COMMUNICATIONS 71

J. RaptorRes. 37(1):71-75 ¸ 2003 The Raptor ResearchFoundation, Inc.

RED-SHOULDEREDHAWK AND APLOMADO FALCON FROM QUATERNARY ASPHALT DEPOSITS IN CUBA

WILLIAM SU•REZ MuseoNacional de Historia Natural, Obispo61, Plaza deArmas; La Habana CP 10100 Cuba

STORRS L. OLSON • NationalMuseum of Natural History,Smithsonian Institution, Washington, DC 20560 U.S.A.

KEYWORDS: AplomadoFalcon; Falco femoralis;Red-shoul- MATERIAL EXAMINED deredHawk; Buteo lineatus;Antilles; Cuba; extinctions; Jbssil Fossils are from the collections of the Museo Nacional birds;Quaternary; West Indies. de Historia Natural, La Habana, Cuba (MNHNCu). Mod- ern comparativeskeletons included specimensof all of The fossil avifauna of Cuba is remarkable for its diver- the speciesof Buteoand Falcoin the National Museum of Natural History, Smithsonian Institution, Washington, sity of raptors,some of very large size, both diurnal and DC (USNM). The following specimenswere used tbr the nocturnal (Arredondo 1976, 1984, Su'•rez and Arredon- tables of measurements: Buteo lineatus 16633-16634, do 1997). This diversitycontinues to increase(e.g., Su'•- 17952-17953, 18798, 18846, 18848, 18965, 19108, 19929, rez and Olson 2001a, b, 2003a) and many additionalspe- 290343, 291174-291175, 291197-291200, 291216, cies are known that await description. Not all of the 291860-291861, 291883, 291886, 296343, 321580, raptors that have disappearedfrom Cuba in the Quater- 343441, 499423, 499626, 499646, 500999-501000, nary are extinct species,however. We report here the first 610743-610744, 614338; Falcofemoralis 30896, 291300, 319446, 622320-622321. recordsfor Cuba of two widespreadliving speciesthat are not known in the Antilles today. Family Accipitridae Thesefossils were obtainedduring recentpaleontolog- Genus ButeoLacepede ical exploration of an asphalt deposit, Las Breasde San Red-shouldered Hawk Buteolineatus (Gmelin) Felipe, which is so far the only "tar pit" site known in (Fig. 1 A-C) the West Indies. Two fossiliferous localities known as San REFERRED ]V[ATERIAL Felipe I and II occur among extinct and activetar seeps Proximal end of right femur (MNHNCu P4614), distal in the floor of the San Felipe Valley, MatanzasProvince, halves of right and left tibiotarsi (MNHNCu P4615, 5.5 km west of the town of Mart• (ca. 22ø57'N, 80ø58'W; MNHNCu P4616), distal end of left tibiotarsus sheetMart/4084-IV, 1:50 000 map, X502, Y347;map pub- (MNHNCu P4617) and distalhalves of right and left tar- lished in 1986 by the Instituto Cubano de Geodesiay sometatarsi (MNHNCu P4618, MNHNCu P4619). Col- Cartogra•a). The age of the depositsis Quaternary,prob- lected in November 1988 by Manuel Iturralde-Vinent, ably late Pleistoceneand early Holocene (Iturralde-Vi- Reinaldo Rqjas-Consuegra,and Stephen Diaz-Franco at nent et al. 1999, 2000). Although the fossil record of San Felipe II. birds in Cuba has hitherto been biasedby the fact that almostall specimenshave come from cavedeposits, the COMPARISONS tar seepsof San Felipe provide a much better sampleof open-countryand aquatic birds that seldom or never are In size and proportions, these specimensagree with preserved in caves.The list of taxa is extensive and in- the Red-shouldered Hawk (Buteolineatus) (Table 1), be- cludesamong other taxa cranes (Grus), thick-knees(Bu- ing larger than the Broad-wingedHawk (B. p&typterus) rhinus), storks (Ciconiidae), waterfowl (Anatidae), crows and smaller than the Red-tailed Hawk (B. jamaicens•s), (Corvus),with a diversevariety of raptorsand scavengers the only two speciesof Buteothat are year-roundresidents being especiallyabundant (Iturralde-Vinent et al. 2000, in Cuba today (Garrido and Garcia Montafia 1975). As Suarez 2000, Su'•rezand Olson 2003a, b, Su'•rezunpubl. wasthe casewith fossilsfrom the Bahamas,we took pmns data). to compare the specimenswith skeletonsof Gray Hawk (B. nitidus),a widespreadspecies of open country that •s of approximatelysimilar size and that might be expected to have occurred in the West Indies. But skeletal ele- •Correspondingauthor's e-mail address:olson.storrs@ ments of B. nitidusare consistentlymore robust than m nmnh.si.edu B. lineatus. 72 SHORT COMMUNICATIONS VOL. 37, NO. 1

Figure 1. A-C, modern Red-shoulderedHawk, Buteolineatus (USNM 17953,on the left in eachpair) comparedwith Cuban fossilsof the stonespecies (A, MNHNCu P4614; B. P4615; C, P4618). D-E, modern Aplomado Falcon, Falco femo•nlis(USNM 291300, on the left in each pair) compared with Cuban fossilsof the same species (D, MNHNCu P4606; B, P4609). A, proximal end of right femur in anterior view; B, distal end of left tibiotarsusin anterior view; C, distal end of right tarsometatarsusin anterior view; D, right carpometacarpiin internal view;E, left tarsometatarsi in anterior view. Scale bars = 2 cm. M_ARCH2003 SItORT COMMUNICATIONS 73

Table 1. Skeletal measurements (ram) of Cuban fossil and modern Red-shouldered Hawk (Buteo lineatus).

CUBAN FOSSILS MODERN

MEASUREMENT RANGE MEAN N RANGE MEAN N

Femur Depth of head 4.9 1 4.5-5.7 5.1 33 Tibiotarsus

Least width of shaft at midpoint 5.6 1 4.8-6.1 5.5 33 Distal width throughcondyles 9.9-10.9 10.4 2 9.5-11.7 10.9 33 Tarsometatarsus

Least width of shaft at midpoint 4.9 I 3.7-5.0 4.4 31 Width of shaft proximal to meta- tarsal facet 5.1-5.3 5.2 3 4.1-5.8 5.1 32 Depth of shaft proximal to meta- tarsal facet 4.2-4.4 4.3 2 3.4-4.4 3.9 32 Distal width 12.3 1 1 1.2-13.3 12.3 32 Depth of middle trochlea 4.9 1 4.4-5.3 4.9 32

REMARI•q ance of thc intermediate-sized Red-shouldered Hawk

Although the Red-shouldered Hawk now has an en- from Cuba could be related to the disappearanceof prey, tirely continental distribution, it has previously been which would presumably have affected the other specms of Buteo as well. known in the West Indies fi-om a few fossils fi-om cave depositsin the Bahamas(Olson 2000), where it first was Ridgway'sHawk (Buteoridgwayi), endemic to Hispan- described as an endemic genus and species Calohierax iola, is now believed to be a small derivative of B. lineatus quadratus(Wetmore 1937, but seeOlson and Hilgartner (Olson 2000). The prehistoric occurrence of the latter 1982, Olson 2000). Thus, its occurrence in Cuba might in Cuba suggeststhat the ancestral stock of Ridgway's have been predicted. The Bahaman population wasprob- Hawk was probably derived from insular populations of ably derivedti-om that of Cuba, as hasbeen the casewith B. lineatus,and most likely t?om Cuba. many other birds (Brodkorb 1959, Olson and Hilgartner Family Falconidae 1982). Genus Falco Linnaeus, 1758 The Red-shoulderedHawk is ordinarily a speciesof Aplomado Falcon FalcofemoralisTemminck, 1822 roesic bottomland forests, so its withdrawal t?om Cuba (Fig. 1 D-E) and the Bahamasis difficult to understand in light of the REFERRED MATERIAL thct that ecological conditions in these islands presumably have become more roesic since the end of the last Right carpometacarpus lacking minor metacarpal glacial period. Potential sourcesof food were much great- (MNHNCu P4606), right carpometacarpuslacking distal er in Cuba than in the Bahamas, making the disappem• end and minor metacarpal(MNHNCu P4607), distalend ance of this hawk from Cuba even more enigmatic. of left tibiotarsus (MNHNCu P4608), proximal end of The Red-tailed Hawk and the Broad-winged Hawk, left tarsometatarsus (MNHNCu P4609), collected 25 Feb- each represented by supposedlyendemic subspecies(B. ruary 2001 by Stephen Diaz-Franco and William Suarez jamaicensissolitudinis Barbour and B. platypteruscubanensis at San Felipe I, area C.

Burns), are common on Cuba today (Rafihele et al. 1998, COMPARISONS Carfido and Kirkconnell 2000) and both have been re- corded from Quaternary cavedeposits on Cuba (Jim6nez These specimensagree perfectly in size and characters 1997, Suftrezand Arredondo 1997), with the latter being with the Aplomado Falcon (Falcofemoralis) (Table 2). found in the San Felipe II asphalt depositsas well (Suftrez They are much too large for American Kestrel, Merhn, unpubl. data.). It hardly seems likely that the disappear- or Bat Falcon (E sparverius,E columbarius,E rufigular•s) 74 SHORT COMMUNICATIONS VOL. 37, NO. 1

Table 2. Skeletal measurements(mm) in fossil and modern Aplomado Falcon (Falcofemoralis).

CUBAN FOSSILS MODERN

MEASUREMENT RANGE MEAN N RANGE MEAN N

Carpometacarpus Total length 41.8 1 37.7-42.5 40.8 4 Proximal width 4.6* 1 4.4-5.1 4.7 4 Proximal depth 10.8 1 9.1-11.6 10.7 4 Width of major metacarpal at midpoint 3.5-3.6 3.5 2 2.9-3.6 3.3 4

Tibiotarsus Distal width 7.5 a 1 7.0-8.7 7.9 5

Tarsometatarsus

Width at level of proximal foramina 6.4 I 5.7-7.0 6.5 5

Estimated. or for the extinct Cuban speciesF. kurochkini(Sufirez and fossil specimens to the Smithsonian Institution from Olson 2001a), and too small for a Peregrine (E peregri- Cuba. Photographs are by John Steiner, Smithsonian nus) or Prairie falcon (E mexicanus).No skeletons were Photographic Services, and the figure was arranged by Brian Schmidt, Division of Birds. avadable for Orange-breasted Falcon (E deiroleucus'),but th•s specieshas very different proportions from E femor- LITERATURE CITED ahs, with a proportionately shorter and much more robust tarsometatarsus. Am•t)ONDO,O. 1976. The great predatorybirds of the Pleistocene of Cubaß Smithson. Contrib. Paleobiol. 27: 169-188. That the Aplomado Falcon once occurred in Cuba is ß 1984. Sinopsisde las aveshalladas en dep6sitos perhaps not unexpected. It is a partially migratory spe- fosillferos pleisto-holoc6nicosde Cuba. Pep. Invest. cies with an extremely wide range extending from the Inst. Zool. 17:1-35. southwesternUnited Statesto Ticrra del Fuego and the BRODKORB,P. ]959. Pleistocene birds from New Provi- Falkland Islands. It inhabits shrub grasslandsand savanna dence Island, Bahamas. Bull. Fla. State Mus. Biol. Sci. and there is increasing evidence of various species of 4:349-371. b•rds adapted to such conditions in the Quaternary of GA•dDO, O.H. ANDE GA•CI^MONrA•^. 1975. Catfilogo Cuba. This is the first indication of the speciesanywhere de las Aves de Cuba. Academia de Ciencias de Cuba, •n the West Indies. Habana, Cuba. --AND A. KIm4CONNELL.2000. Field guide to the RFSUMEN.--Procedentesde dep0sitoscuaternarios de as- birds of Cuba. Cornell Univ. Press, Ithaca, NY U.S.A. falto en San Felipe, al norte de la Provinciade Matanzas, ITURRAI•DE-VINENT, M., R.D.E. MACPHEE, S. DiAZ-FRANCO, se registran por primera vez para Cuba dos especiesde AND R. ROJAS-CONSUEGRA.1999. A small "Rancho La rapaces que viven hoy en el continente: Buteolineatus y Brea" site discoveredin Cuba.J. Geol.Soc. Jam. 33:20. Falco]bmoralis;este filtimo constituyela primera evidencia , --, --, W. SU•REZ, AND A. LOM- de cse taxon cn la Subregi6 n Antiliana. BA.'2000. Las Breas deSan Felipe, aQuaternary as- [Traducci6n de los autores] phalt seep near Marti (MatanzasProvince, Cuba). Ca- ribb.J. Sci.36:300-313. ACKNOWLEDGMENTS JIMP.NEZ VASQtJEz, O. 1997. Seisnucvos registros de aves Travel by W. Sufirez to Washington,DC, wasmade pos- f6siles en Cuba. Pitirre 10:49. sible by the Alexander Wetmore endowment fund, Divi- OLSON, S.L. 2000. Fossil Red-shouldered Hawk in the Ba- sion of Birds, National Museum of Natural History, hamas: Calohieraxquadratus Wetmore synonymized Sm•thsonian Institution. W. Su•trez is especiallygrateful to Dr. Helen James (Smithsonian Institution), and Travis with Buteo lineatus (Gmelin). Proc. Biol. Soc.Wash. 113: and SydneyOlson, for their help and kindnessduring 298-301. h•s visit to Washington, DC. Gilberto Silva Taboada (Mu- --AND W.B. HILGARTNER. 1982. Fossil and subfossil seo Nacional de Historia Natural de Cuba) transported birds f•om the Bahamas. Pages 22-56 in S.L. Olson MARCH 2003 SHORT COMMUNICATIONS 75

[ED.], Fossil vertebrates from the Bahamas. Smithson. Caracaracreightoni Brodkorb based on fossilsfi'om the Contrib. Paleobiol. 48. Quaternary of Cuba (Aves:Falconidae). Proc.Biol. Soc RAFFAELE,H.A., J. WILEY,O. GARR1DO,A. KEITH,AND J. Wash. 114:501-508. RAFFAELE.1998. A guide to the birds of the West In- -- ANt)--. 2003a. A new speciesof caracara(Mzl- dies. Princeton Univ. Press,Princeton, NJ. vago)from Quaternary asphaltdeposits in Cuba, w•th SUAREZ, W. 2000. Gontribuci6n al conocimiento del es- notes on new material of Caracaracreightoni Brodkorb tatus genarico del c6ndor extinto (Giconiiformes: (Aves:Falconidae). Proc. Biol. Soc. Wash. 16: in press Vulturidae) del Guaternario cubano. Ornitol.Neotrop. --AND --.. 2003b. New records of storks (Cico- 11:109-122. niidae) from Quaternary asphalt deposits in Cuba --AND O. ARREDONDO. 1997. Nuevas adiciones a la Condor 105:150-154 paleornitologia cubana. Pitirre 10:100-102. WETMORE,A. 1937. Bird remainsfrom cavedeposits on -- ANDS.L. OLSON.2001a. A remarkablenew species Great Exuma Island in the Bahamas.Bull. Mus. Comp. of small falcon from the Quaternary of Guba (Aves: Zool. 80:427-441. Falconidae: Falco). Proc. Biol. Soc. Wash. 114:34-41. --AND --'. 200lb. Further characterization of Received22 July 2002; accepted28 November 2002

j. RaptorRes. 37 (1) :75-77 ¸ 2003 The Raptor ResearchFoundation, Inc.

SUBADULT AND PALE STEPPE EAGLES BREEDING IN MONGOLIA

DAVID H. ELLIS 1 USGSPatuxent Wildlife Research Cente•; 11410 AmericanHolly Drive, Laurel, MD 20708 U.S.A.

KEYWORDS: SteppeEagle;, Aquila nipalensis. placement (i.e., newly grown) feathers are light or dark before concludingthat the Steppe Eagle has a pale adult All adult Steppe Eagles (Aquila nipalensis)are report- morph. Although subadultsof some speciesof Aquila ea- edly very dark (Ferguson-Leesand Christie 2001). How- gles are known to at least occasionallybreed (e.g., New- ever, the closely related (Wink and Sauer-Gfirth 2000) ton 1979,Steenhofet al. 1983), I knowof no prior record Tawny Eagle (A. rapax) does have a pale adult morph of a subadultSteppe Eagle breeding. (see Plate 114, Brown and Areadon 1968). Clark (1992) During five expeditionsto Mongolia from 1994-2000, decried the confusion in the scientific literature and in I found more than 20 Steppe Eagle nests.At one site •n museum collections over the various morphs of the arid southeasternMongolia (115øE, 45øN), we found a Steppe and Tawny eagles and advanced "criteria for the very pale bird (Fig. 1). Elsewherewe found two rufous- correct identification of all museum specimensand live plumagedbirds. All three were attending live young. One birds .... "He statesconclusively that SteppeEagles be- of the rufous birds was captured (Ellis et al. 2001) and come "much darker as adults." His assertion stems from photographed in hand. The very pale bird was photo- fieldwork in Israel, India, and Africa, and, more impor- graphed on its nest at a distance of 2 m. tanfly, from handling over 300 museum specimens.His In Mongolia, Steppe Eagle adults are generally deep conclusionreaffirms statementsby Cramp and Simmons chocolatebrown above and below with blackishremiges (1980) that subadultsare paler than adults and that all and rectrices finely barred with black. These dark birds very pale birds are young. match Glark's (1992, 1996) descriptions of the dark While it is helpful to examine migrantsand wintering brown adult plumage. The only consistentlypresent light birds in evaluating the prevalenceof adult morphs, evi- area in the plumage of dark adults is a broken line of dence to support the claim that no adults are pale must whitish spotson the upper tail coverts.This line is readdy come from the breeding grounds.Even there, if very pale visible at great distanceswhen a bird is flying, except breeders are found, it is necessaryto determine if re- when overhead.Some dark birds (probably thosetoolung from subadult plumage) show a dappled line of light brown at the trailing edge of the under wing coverts. l Present address:USGS SouthwestBiological Science Under some light conditions, pale areas at the base of Center, HC 1 Box 4420, Oracle, AZ 85623 U.S.A.; e-mail the primaries are apparent on the underside of the wings address: dcellis@theriver. com of some, and probably all, dark adults. The head is gen-