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A New Species Of Owl Of The Genus Bubo From The Pleistocene Of Cuba PDF

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Preview A New Species Of Owl Of The Genus Bubo From The Pleistocene Of Cuba

PROC. BIOL. SOC. WASH. 107(3), 1994, pp. 436-444 A NEW SPECIES OF OWL OF THE GENUS BUBO FROM THE PLEISTOCENE OF CUBA (AVES: STRIGIFORMES) Oscar Arredondo and Storrs L. Olson Abstract.—Bubo osvaldoi, new species, is described from three bones from aPleistocenecavedepositintheSierradeGaleras,PinardelRio,westernCuba, with two additional paratypes referred from Baire in the province ofSantiago deCuba, inthe eastern part ofthe island. The speciesis the onlyrepresentative ofthe genus Bubo and the tribe Bubonini in the Antilles and in size was larger than any living owl. Resumen.—Se describe una especie nueva de buho, Bubo osvaldoi, de tres huesos del Pleistoceno hallados en una cueva de la Sierra de Galeras, en Pinar del Rio, Cuba occidental, yde otrosdoshuesosparatiposprocedentesde Baire, provincia de Santiago de Cuba, en el oriente de la isla. La especie es el unico miembro del genero Bubo y de la tribu Bubonini conocido hasta ahora en las Antillasyeramayorquecualquieradelasformasconocidasdebuhosvivientes. The Quaternary avifauna ofCuba is re- western end of Cuba, along with a manu- markable for the number and size of the script describing them as a new species of extinctowlsthathavebeendiscoveredthere Bubo. This called to mind two enigmatic (summarized by Arredondo 1976, 1982). bones that Olson had studied some years These include the truly enormous strigids previouslythatcamefromtheformerprov- of the endemic genus Ornimegalonyx, of ince ofOriente, at the opposite end ofthe which four species have been named (Ar- island. These had originally been sent in redondo 1982),andtwospeciesofbamowls January 1947 byAbelardo Moreno, then of {Tyto) that farexceed in size anyliving spe- the Museo Poey, to AlexanderWetmore, at cies of Tytonidae. Pulsatrix arredondoi the Smithsonian Institution. Wetmore had (Brodkorb 1969) is slightly smaller than its had a drawing made ofthese bones and left NeotropicalrelativeP.perspicillata, butbe- afolderofcorrespondenceconcerningthem, longs to a genus otherwise unknown in the but they remained unpublished. We have Antilles. Itwaspreviouslyknownonlyfrom concludedthatthese probablybelongto the the type locality, Cueva Paredones in Ha- same species as represented by the bones bana Province, but Arredondo has recently from Pinar del Rio, which cannot be re- identified a tarsometatarsus ofP. arredon- ferred to any known species of owl, living doi from Cueva Calero in Matanzas Prov- or fossil. ince. Here we present evidence ofyet another Class Aves Linnaeus, 1758 very large extinct Cuban owl, amidst what Order Strigiformes Wagler, 1830 couldalreadyhavebeenregardedasapleth- Family Strigidae Vigors, 1825 ora. Our collaboration on this began when Arredondo forwarded for evaluation three The fossil material treated below may be fossil specimens from Pinar del Rio, at the distinguished from Tyto (Tytonidae) by the VOLUME 107, NUMBER 3 437 Fig. 1. Tarsometatarsi ofstrigid owls. A, anterior view of Ornimegalonyx oteroi (MNHN unnumbered, lackingdistalend),showingtheimmensesizecomparedtootherwiseverylargeowls;B,C,D,posterior,anterior, and distal views ofBubo osvaldoi, new species, holotype (MNHNCu-27, coated with ammonium chloride to enhancedetail); E, anteriorviewofB. bubo(USNM 610384) female, largestavailablespecimen ofthe species. Figuresthree-fourthsnatural size. proportionately short, robust tarsometatar- gigantic Cuban owls ofthe genus Ornime- sus, the more anteroposteriorly expanded galonyx, in which the proportions of the external condyle ofthe femur, and the shal- tarsometatarsus are very different (Fig. 1). lower brachial depression ofthe humerus. They are clearly referable to the group of largest strigid owls (Tribe Bubonini) that includes the genera Bubo, Nyctea, Ketupa, Genus Bubo Dumeril, 806 1 and Scotopelia, concerning which Ford The Cuban fossils, although from a very (1967:82) wrote: "Each ofthese genera has large owl, are nonetheless dwarfed by the some osteological characteristics by which 438 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON it can be separated, but the diJSerences are Measurementsofholotype(mm).—Length notgreatanditisquestionableastowhether from distalend ofattachment ofM. tibialis or not they actually warrant separation." anticus to trochleae, 63.3; least width of Wewouldechothesesentiments,aswelike- shaft, 11.9; depth ofshaft at midpoint, 7.8; wise find little in the osteology of at least distal width across trochleae, 24.8; depths Nyctea or Ketupa that would merit sepa- ofinner, middle, and outertrochleae, 12.1, ratingthemfromBubo(wedidnotexamine 9.8, 14.9. skeletons ofScotopelia). Ketupa and Scoto- Topotypical paratypes.—Corax>^e\t right pelia are Asian and African genera, respec- femur, lacking the internal condyle and MNHNCu tively, whose traditional generic characters abraded about the trochanter, involve external aspects ofthe feet that are 27.3 (Fig. 2B). Shaftoflefttarsometatarsus, adaptations for fishing. Nyctea is hardly lackingtheproximalportionandthedigital more than a Bubo adapted to life in Arctic trochleae, MNHNCu-27.2. These speci- tundra. On geographical grounds alone. mens and the holotype appear to be very Bubo is the only genus of Bubonini likely heavilymineralizedandareblackishincol- to have colonized Cuba, and we find noth- or, mottled with orangish brown. The par- ing in the Cuban fossils to suggest a closer atypicaltarsometatarsusisfromanindivid- relationshipto any ofthe otherowls ofthat ual slightly larger than represented by the tribe than to Bubo. In Strix nebulosa, the holotype and the specimen is highly pol- largest New World member ofthe Strigini, ished and evidently water worn. the femurandhumerasare proportionately Additionalparatypes.—Rightfemurlack- much more elongate and gracile, the wing inginternalcondyle,anteriorsurfaceofhead, oftheinnertrochleaislesspronounced, and and a piece out ofthe posterior face ofthe theattachmentofM.tibialisanticusismore shaft,USNM447022(Figs. 2A, 3A-C).Left distinctandraisedthanintheCubanfossils, humerus lacking proximal end and the ex- which conform with Bubo in all these re- ternalpartofthedistalarticulation, USNM spects. 447023 (Figs. 3D, 4B). The only informa- tion we have concerningthese specimens is that they came from a "mine" (perhaps a Bubo osvaldoi, new species cavern exposed in roadwork or similar ac- Figs. 1-4 tivity)inthevicinityofBaire,OrienteProv- Holotype.—Kight tarsometatarsus of an ince (now in the Provincia de Santiago de adult lacking the proximal end above the Cuba). They are creamy white in color and attachment for M. tibialis anticus (Fig. IB, areverylighttothetouch,asthoughlacking C). Deposited in the Museo Nacional de organic constituents, perhaps through Historia Natural de Habana (MNHNCu- leaching. The same site is the type locality 27.1). of the sloth Neocnus baireiensis Mayo Type locality.—Cueva del Mono Fosil, (1980),describedfrommaterialpresumably southern side ofthe Sierra de Galeras, Cor- collected at the same time as the owl bones dillera de Guaniguanico, Municipality of andforwhichthereislikewisenomorepre- Vinales, Province of Pinar del Rio, Cuba. cise information. This is also the type locality of the fossil Measurements ofparatypes (mm).—Tar- platyrrhine monkey Paralouatta varonai sometatarsus MNHNCu-27.2: least width (Rivero and Arredondo, 1991). of shaft, 1 1.5; depth of shaft at midpoint, ^^e.—Quaternary, probably Pleistocene, 7.2 (worn). Femur MNHNCu-27.3: length, at least asjudgedby the mineralization and 101.5; proximal width 22.7; depth ofhead associated fauna ofthe holotype and topo- 9.4; width and depth ofshaft at midpoint, types. 9.8 X 9.4; depth through external condyle, VOLUME 107, NUMBER 3 439 Fig. 2. FemoraofBubo in anteriorview. A, B. osvaldoi, newspecies(USNM 447022); B, B. osvaldoi, new species(MNHN27.3,coatedwithammoniumchloridetoenhancedetail);C, B. bubo(USNM610384)female, largestavailablespecimenofthespecies; D, B. bubo(USNM 343007)male; E, B. virginianus(USNM 501314) female; F, B. virginianus (USNM 555903), male. Size differences between the two femora offi. osvaldoi are closely comparable to those between sexes in other species ofthe genus. Note the different position ofthe intermuscularline inB. osvaldoi. Figuresthree-fourthsnaturalsize. 19.0. FemurUSNM 447022: length, 1 12.4; tarsometatarsus much larger. The inter- proximal width 27.7; depth of head 10.3; muscular line on the anterior face of the width and depth ofshaft at midpoint, 11.5 shaft ofthe femur is unique in originating X 9.9; depththroughexternalcondyle, 21.9. at the distal end of the trochanteric crest Humerus USNM 447023: length from dis- and running diagonally across the shaft, tal end of pectoral crest to entepicondyle, whereas in Bubo and other owls examined 99.1; length (diagonal) ofbrachial depres- the line originates at the neck ofthe femur sion, 17.6. or near the trochanter and runs parallel, or Etymology.—The species is dedicated to nearly so, to the sides ofthe shaft (Fig. 2). the discoverer ofthe specimens from Pinar Humerus small relative to the hindlimb. del Rio, Osvaldo Jimenez, speleologist and Comparative material examined.—Skel- amateur paleontologist. etons from the collections of the National Diagnosis.—Largerthananymodemspe- Museum ofNatural History, Smithsonian cies ofBubonini (Table 1), which includes Institution (USNM) as follows: Bubo bubo the largest modem owls. Distal foramen of (4 males, 1 female, 1 unsexed, 1 female 440 PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON B Fig. 3. DrawingsofthetwoparatypesofBuboosvaldoi, newspecies,fromBaire,SantiagodeCuba:A,femur inposteriorview(USNM447022);B,same,anteriorview;C,same,proximalview; D,humerusinpalmarview (USNM 447023). Natural size. trunk; Sweden, Greece, and Korea), Bubo and humeri of a female; Kenya and Zim- virginianus (6 males and 6 females; Penn- babwe), Bubo philippensis (1 female; Lu- sylvania, District of Columbia, Florida, zon), Bubosumatrana (1 unsexed; Borneo), Minnesota, NewMexico, Arizona, andCal- Ketupaketupu(1 femaleand 1 femaletrunk; ifornia),Buboafricanus(2 males, 4 females, Borneo and Java), Ketupa blakistoni (1 un- and 1 male trunk; Sudan, Tanzania, Zim- sexed; Japan), Ketupa zeylonensis (2 fe- babwe, andTransvaal), Bubolacteus{\ male males and 1 unsexed; Thailand and un- VOLUME 107, NUMBER 3 441 Fig. 4. HumeriofBuboinpalmarview. A, B. bubo (USNM 610384)female, largestavailable specimen of the species; B, B. osvaldoi, new species (USNM 447023); C, B. virginianus (L[SNM 501314), female. Figures three-fourthsnaturalsize. knownlocalities), Nycteascandiaca(2 males Chauvire &Weesie (1986), mostoftheoth- and 2 females; Alaska, Maryland, and Dis- erfossilspeciesattributedtothegenusBubo trictofColumbia).Thefollowingspecimens are smaller than B. insularis. According to ofScotopelia in the American Museum of Brodkorb&Mourer-Chauvire(1984),Bubo Natural History were measured and ex- Sinclair! L. Miller, based on scanty and amined for us: S. peli(2 unsexed; Zaire), S. poorly preserved material from Quaternary bouvieri(1 unsexed; Zaire), S. ussheri (1 fe- cavesitesinCalifornia, fallswithintherange male; Sierra Leone). of size variation of B. virginianus and is Comparisonwithotherfossilspecies.—The probably a synonym ofthat species. only otherextinct insular species ofBubois BuboflorianaeKretzoi, 1958, isbased on B. insularis from Corsica and Sardinia, a pedal phalanx from the early Pliocene of which was described as being smaller than Hungarythatprobablyfallswithintherange B. bubo(Mourer-Chauvire&Weesie 1986), of size variation of Bubo bubo (Janossy and hence is even smaller than B. osvaldoi. 1977). Bubo bubodavidi(Mourer-Chauvire As ascertained from the review in Mourer- 1975) was described from the Pleistocene 442 PROCEEDINGSOFTHEBIOLOGICALSOCIETY OFWASHINGTON Table L—Selected measurements oflarge owls (Strigidae) compared with Bubo osvaldoi, new species. For numbersofspecimensseeComparativeMaterialExamined.FEM=lengthoffemur.HUM=lengthofhumerus from distal end of pectoral crest to entepicondyle. TMT 1 = length oftarsometatarsus from distal end of attachmentofM. tibialis anticusto trochleae. TMT 2 = widthoftarsometatarsusacrosstrochleae. Species FEM HUM TMT 1 TMT2 Buboosvaldoi 101.5-112.4 99.1 63.3 24.8 Bubobubo 93.5-104.5 97.7-110.7 46.6-50.4 18.9-23.0 Bubo virginianus 75.1-86.1 76.9-89.7 36.5-41.0 16.1-19.2 Buboafricanus 64.5-72.2 69.6-74.6 42.0-52.4 13.1-13.9 Bubolacteus 91.1 104.9-106.4 46.4 23.1 Bubophilippensis 77.2 85.0 43.0 20.8 Bubosumatrana 60.6 75.3 28.2 17.5 Ketupa ketupu 75.9-78.9 81.8 51.1 15.9 Ketupa blakistoni 102.7 116.7 48.7 21.2 Ketupa zeylonensis 82.0-97.1 87.3-104.1 52.4-56.1 16.3-19.9 Nycteascandiaca 83.8-92.6 92.8-102.4 28.3-32.0 17.6-20.2 — — Scotopeliapeli 92.1-96.2 22.2 — — Scotopelia bouvieri 65.3 13.2 — — Scotopelia ussheri 73.8 14.2 ofFrance as a larger temporal form ofthe Homed Owl {Bubo virginianus), which has modem species, although there is overlap, probably the greatest latitudinal breeding particularly with the largest specimen of rangeofanybirdintheworld, nestingfrom Bubobubothatweexamined,whichexceeds northern Canada and Alaska to Tierra del any ofthose reported by Mourer-Chauvire Fuego. Itdoesnot occurin the West Indies, (1975). Therewereotherwisenoqualitative however, and Bubo osvaldoiis thusthe first differences shown by the fossil form and it and only member of the Bubonini recog- cannot, therefore, beidentifiedwiththeCu- nized in the Antilles. ban bird. In size. Bubo osvaldoi exceeds any living BubobinagadensisBurchak-Abramovich, owl (Table 1), there beingno overlapintar- 1965, from the Pleistocene of Azerbaijan, sal measurements. The smaller ofthe two was described from a femur that was said fossil femora is exceeded only by the one to be not less than the size of that in the unsexed specimen ofKetupa blakistoniand eastern forms offishing owls (genus Ketupa by the largest female specimen available of assumed, although we may wonder ifBur- Bubo bubo. As females are the larger sex in chak had access to any comparative mate- owls, ifwe assume that the small fossil fe- rial) and much larger than in Bubo bubo. mur is from a male, then on a sex per sex Neither illustrations nor measurements ac- basis there would probably be no overlap companied the description, only some in- in any hindlimb measurements between decipherable "indices of massiveness," so Buboosvaldoiandlivingowls.Thehumerus wecannotdetermineanythingfurtherabout inB. osvaldoi, however, fallswithinthe size this taxon at present. range ofseveral ofthe larger species ofowls — Discussion. Mxhough bubonine owls are (Table 1), suggesting that the wing in this diverse in the Old World, only two taxa insular form may have been reduced rela- occur today in the New World—the Snowy tive to the overall size ofthe bird. Owl {Nyctea scandiaca), which is circum- There is considerable variation in pro- polar in the high Arctic, and the Great portions ofthe hindlimb, especiallythetar- VOLUME 107, NUMBER 3 443 from a lineage otherthanthatgiving rise to B. virginianus may yet be found. Acknowledgments We are grateful to Ross MacPhee for transporting specimens and papers from Cuba, to Orlando H. Garrido for returning same, and to Allison V. Andors for supply- ing measurements and notes on skeletons ofScotopelia in the American Museum of Natural History, New York. Prior to the discoveryofthetypematerial,WilliamHil- gartner made valuable comparisons ofthe specimens from Oriente confirming that these were not tytonine and probably rep- We resented a species ofBubo. also thank theGrupoEspeleologico"PedroA. Borras" oftheSociedadEspeleologicadeCubamak- Fig. 5. Tarsometatari ofBuboin anterior view to ingthe owl fossils foundby one ofits mem- showthegreatdifferencesinproportionsbetweenvar- bers in Pinar del Rio available for study. iousspecies. A,B. virginianus(USNM 613846); B,B. The photographs are by Victor E. Krantz africanus(USNM490288); C,B. sumatranus(USNM and the drawings by L. Isham. 559827). Figuresthree-fourthsnatural size. Literature Cited sometatarsus, in Bubo. Compared to B. Arredondo,O. 1976. Thegreatpredatorybirdsofthe bubo, the tarsometatarsus in B. africanus, Pleistocene ofCuba [Translated and amended for example, is quite long and slender, byS.L.Olson].Pp. 169-187inS.L.Olson,ed., whereas that in B. sumatranus is extraor- SmithsonianContributionstoPaleobiology27. dinarily short and stout (Fig. 5). That ofB. c.u1b9a8n2o..—LoBoslSettriingdiefolramSesocfioseidlaedsdVeelnpelzeoilsatnocaednee osvaldoi is ofmore typical proportions and CienciasNaturales 140:33-55. is similar to that ofB. bubo. Brodkorb, P. 1969. AnextinctPleistoceneowl from It is not necessary to assume that B. os- Cuba.—QuarterlyJournalofthe FloridaAcad- avanludsoisiismpmloystbecclaousesleytrheelaltaetdtetroiBs.thveirgoinnliy- ef,rm&oymCo.efaSrMlcoyiuermnecaren-sC3hs1iat:ue1vs1i2or-fe1.O1l4d.1u9v8a4i. GoForsgsei,l Toawnl-s geographically proximate bubonine in the zania.—Ostrich 54:17-27. New World. Olson (1984) reported a man- Burchak-Abramovich, N. L 1965. Noviyavid isko- dibular symphysis ofa very large owl from paemogo filina iz Binagadov.—Ornitologiya thePleistoceneatLadds, Georgia,thatsure- Ford, N7.:4L5.2-415946.7.[InARussyssitaenm]atic study ofthe owls ly represents a species otherwise as yet un- based on comparative osteology. Unpublished known in North America. Although this Ph.D.dissertation.UniversityofMichigan,Ann specimen seemed to be more similar in Arbor. University Microfilms 68-7595. morphology to Strixthan to Bubo, it is not Janossy,D. 1977. Plio-Pleistocenebirdremainsfrom verydiagnostic. Regardless,itindicatesthat theCarpathian Basin IIL Strigiformes, Falcon- iformes, Caprimulgiformes, Apodiformes.— extinct lineages oflarge owls remain to be Aquila 84:9-36. discovered in the fossil record of North Kretzoi, M. 1958. Birdremainsfromthe Hipparion America, so that a progenitor of.6. osvaldoi fauna ofCsakvar.—Aquila 63-64:239-248. 444 PROCEEDINGSOFTHE BIOLOGICALSOCIETYOFWASHINGTON Mayo, N. A. 1980. Nueva especie de Neocnus (Ed- Georgia. Pp. 44—46 in H. H. Genoways & M. entata: Megalonychidae de Cuba) y considera- R. Dawson, eds., Contributions in Quaternary cionessobrelaevolucion, edadypaleoecologia vertebratepaleontology:Avolumeinmemorial de las especies de este genero.—Actas del Se- toJohnE.Guilday.—CarnegieMuseumofNat- gundo Congreso Argentine de Paleontologia y ural History Special Publication 8. Bioestratigrafia y Primer Congreso Latinoam- Rivero[delaCalle],M.,&O.Arredondo. 1991. Par- ericanode Paleontologia. Volume 3:223-236. alouattavaronai, anewQuaternaryplatyrrhine Mourer-Chauvire, C. 1975. Les oiseaux du Pleisto- from Cuba.—Journal ofHuman Evolution 21: cene Moyen et Superieur de France.—Docu- 1-11. ments des Laboratoires de Geologic de la Fa- cultedes Sciencesde Lyon 64(2):1-624. (OA) Museo Nacional de Historia Nat- , & P. D. M. Weesie. 1986. Bubo insularisn. ural, CapitolioNacional, LaHabana, Cuba; sp., forme endemique insulaire de grand-due (Aves,Strigiformes)duPleistocenedeSardaigne (SLO) Department of Vertebrate Zoology, et de Corse.—Revue de Paleobiologie 5:197- National Museum of Natural History, 205. Smithsonian Institution, Washington, D.C. Olson,S. L. 1984. Averylargeenigmaticowl(Aves: 20560, U.S.A. Strigidae) from the Late Pleistocene at Ladds,

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