Independent geographic origins of the genus Amazona in the West Indies

Main Article Content

Patricia Ottens-Wainright
Kenneth M. Halanych
Jessica R. Eberhard
Rachel I. Burke
James W. Wiley
Rosemarie S. Gnam
Xiomara Gálvez Aqualera

Abstract: Nine species of the parrot genus Amazona are endemic to the Greater Antilles, Bahamas, and Cayman Islands (A. leucocephala, A. agilis, A. collaria, A. ventralis, A. vittata) and Lesser Antilles (A. guildingii, A. imperialis, A. arausiaca, A. versicolor). Populations of one species, A. leucocephala, colonized Cuba, Bahamas, and Cayman Islands resulting in five subspecies. Biogeographic relationships of these Antillean Amazona were examined by a reconstruction of their evolutionary history: mitochondrial cytochrome b sequence data were analyzed with maximum likelihood, parsimony, and distance methods. Phylogenetic analyses show a distinct divergence of the smaller and mostly green Greater Antillean Amazona from the larger, more colorful Lesser Antillean species, and imply that they colonized the West Indies independently. This phylogenetic reconstruction was used to trace potential dispersal routes of ancestral Amazona into the West Indies. The species distribution found today in the Lesser Antilles may have been the result of at least two colonization events from South America, one or more of which occurred early in the history of this genus. Data from this study also suggest that there may have been two dispersal events to the Greater Antilles. The Greater Antillean species appear closely related to the small A. albifrons of Central America. Evolutionary relationships within the A. leucocephala subspecies complex suggest that A. l. bahamensis and A. l. caymanensis were the first populations of this species to become genetically isolated. Isolation of populations on Cuba (A. l. palmarum and A. l. leucocephala) occurred later.


Keywords: Amazona, biogeography, Caribbean, cytochrome b, parrots, phylogeny


Resumen:Los orígenes geográficos independientes del género Amazona en las Antillas- Nueve especies de cotorras del género Amazona son endémicas en las Antillas Mayores, Bahamas y las islas Caimán (A. leucocephala, A. agilis, A. collaria, A. ventralis, A. vittata) y las Antillas Menores (A. guildingii, A. imperialis, A. arausiaca, A. versicolor). Poblaciones de una especie, A. leucocephala, colonizaron Cuba, Bahamas y las islas Caimánn resultando en cinco subespecies. Las relaciones biogeográficas de las especies de Amazona de las Antillas fueron examinadas por medio de una reconstrucción de su historia evolutiva: secuencias de citocromo b mitocondrial fueron analizadas utilizando métodos de parsimonia, máxima verosimilitud y de distancia. Análisis filogenéticos muestran una marcada divergencia entre las especies de las Antillas Mayores, que tienen menor tamaño corporal y plumaje predominantemente verde, y las especies de las Antillas Menores, que tienen plumajes más coloridos. Esta divergencia implica que los dos grupos colonizaron los Antilles independientemente. Esta reconstrucción filogenética fue utilizada para trazar rutas potenciales de dispersión de las Amazona ancestrales por las Antillas. La presente distribución de especies en las Antillas Menores podría ser resultado de por lo menos dos colonizaciones desde Sudamérica, y por lo menos una de éstas ocurrió temprano en la historia del género. Datos de este estudio también sugieren la posibilidad de dos eventos de dispersión a las Antillas Mayores. Las especies de las Antillas Mayores están estrechamente relacionadas con A. albifrons, una especie relativamente pequeña de Centroamérica. Relaciones evolutivas dentro del complejo de A. leucocephala sugieren que A. l. bahamensis y A. l. caymanensis fueron las primeras poblaciones de esta especie en aislarse genéticamente. El aislamiento de las poblaciones de Cuba (A. l. palmarum y A. l. leucocephala) ocurrió más tarde.


Palabras clave: Amazona, biogeografía, Caribe, citocromo b, cotorras, filogenia


Résumé: Origines géographiques indépendantes chez le genre Amazona dans les Antilles- Neuf espèces de perroquets du genre Amazona sont endémiques des Grandes Antilles, des Bahamas et des îles Cayman (A. leucocephala, A. agilis, A. collaria, A. ventralis, A. vittata) et des Petites Antilles (A. guildingii, A. imperialis, A. arausiaca, A. versicolor). Des populations d'une espèce, A. leucocephala, ont colonisé Cuba, les Bahamas et les îles Cayman, donnant naissance à cinq sous-espèces. Les relations biogéographiques entre ces amazones antillaises ont été étudiées par la reconstruction de leur histoire évolutive. Les données de séquences du cytochrome b mitochondrial ont été étudiées par les méthodes du maximum de vraisemblance, de parcimonie et de distance. Les analyses phylogénétiques montrent une divergence entre les amazones des Grandes Antilles, plus petites et à dominante verte, et les espèces plus grandes et colorées des Petites Antilles, ce qui indiquent qu'elles ont colonisé les Antilles indépendamment. Cette reconstruction phylogénétique a été utilisée pour tracer les routes potentielles de dispersion des amazones ancestrales dans les Antilles. La distribution des espèces observées aujourd'hui dans les Antilles pourrait être le résultat d'au moins deux événements de colonisation depuis l'Amérique du Sud, l’un d'entre eux, ou plusieurs, étant survenu très précocement dans l'histoire du genre. Les données de cette étude suggè- rent qu'il pourrait y avoir eu aussi deux événements de dispersion dans Les Grandes Antilles. Les espèces des Grandes Antilles apparaissent étroitement reliées au petit A. albifrons d'Amérique Centrale. Les relations évolutives du complexe de sous-espèces de A. leucocephala suggèrent que A. l. bahamensis et A. l. caymanensis ont été les premières populations de cette espèce à avoir été génétiquement isolées. L?isolement des populations de Cuba (A. l. palmarum et A. l. leucocephala) est survenue plus tard.


Mots clés: Amazona, biogéographie, Caraïbe, cytochrome b, perroquets, phylogénie

Abstract 1403 | PDF Downloads 752

References

ANDERSON, S., A. T. BANKIER, B. G. BARRELL, M.
H. L. DE BRUIJN, A. R. COULSON, J. DROUIN, I.
C. EPERON, D. P. NIERLICH, B. A. ROE, F.
SANGER, P. H. SCHREIER, A. J. H. SMITH, R. STADEN, AND I. G. YOUNG. 1981. Sequence and
organization of the human mitochondrial genome.
Nature 290:457?465.

ARCTANDER, P. 1988. Comparative studies of avian
DNA by restriction fragment length polymorphism
analysis: convenient procedures based on
blood samples from live birds. J. Ornithol.
129:205?215.

ARLDT, T. 1936. Esquisse de géographie zoologique
des Antilles (1). Rev. Soc. Hist. Géogr.
Haiti 8(25):28?40.

ARREDONDO, O. 1984. Sinopsis de las aves halladas
en depósitos fosilíferos Pleisto-holocénicos de
Cuba. Rep. Invest. Inst. Zool., Acad. Cienc. Cuba
17:1?35.

AVISE, J. C. AND D. WALKER. 1998. Pleistocene
phylogeographic effects on avian populations and
the speciation process. Proc. R. Soc. Lond., Ser.
B. 265:457-463.

AVISE, J. C. 2000. Phylogeography: the history and
information of species. Cambridge, MA: Harvard
Univ. Press.

BECKER, J. J. 1987. Neogene avian localities of
North America. Pp. 34?35. Washington, DC:
Smithson. Inst. Press.

BERLIOZ, J. 1959a. Un exemple des particularités de
la faune antillaise: les colibris. C. R. Soc.
Biogéogr. 36(311):3?6.

BERLIOZ, J. 1959b. Le peuplement animal des Antilles:
les oiseaux des Grandes Antilles. C. R.
Soc. Biogéogr. 36(319):115?121.

BERMINGHAM, E., AND H. A. LESSIOS. 1993. Rate
variation of protein and mitochondrial DNA evolution
as revealed by sea urchins separated by the
Isthmus of Panama. Proc. Natl. Acad. Sci. USA
90:2734?2738.

BIRT, T. P., V. L. FRIESEN, J. M. GREEN, W. A.
MONTEVECCHI, AND W. S. DAVIDSON. 1992.
Cytochrome b sequence variation among parrots.
Hereditas 117:67-72.

BOND, J. 1963. Derivation of the Antillean avifauna.
Proc. Acad. Nat. Sci. Phila. 115:79?98.

BOND, J. 1979. Derivations of Lesser Antillean
birds. Proc. Acad. Nat. Sci. Phila. 131:89?103.

BRISSON, M. J. 1760. Ornithologie ou méthode contenant
la division des oiseaux en ordres, sections,
genres, espèces & leurs variétés. Paris: J.-B.
Bauche.

BRODKORB, P. 1959. Pleistocene birds from New
Providence Island, Bahamas. Bull. Fla. St. Mus. Biol. Sci. 4(11):349?371.

BRODKORB, P. 1971. Catalogue of fossil birds: Part
4 (Columbiformes through Piciformes). Bull. Fla.
St. Mus. Biol. Sci. 15(4):163?266.

BUDEN, D. W., AND S. L. OLSON. 1989. The avifauna
of the Cayerias of southern Cuba, with the
ornithological results of the Paul Bartsch Expedition
of 1930. Smithson. Contrib. Zool. 477.

BUSKIRK, R. E. 1985. Zoogeographic patterns and
the tectonic history of Jamaica and the northern
Caribbean. J. Biogeog. 12:445?461.

BUTLER, P. J. 1992. Parrots, pressures, people, and
pride. Pp. 25?46 in New World parrots in crisis:
solutions from conservation biology (Beissinger,
S. R., and N. F. R. Snyder, eds.). Washington,
DC: Smithson. Inst. Press.

CARRAWAY, C., AND P. CARRWAY. 1979. The Bahama
Parrot: Amazona leucocephala bahamensis.
Avic. Mag. 85:18?23.

CAMPBELL, K. E., JR. 1976. The late Pleistocene
avifauna of La Carolina, southwestern Ecuador.
Pp. 155?168 in Collected papers in avian paleontology
honoring the 90th birthday of Alexander
Wetmore (Olson, S. L., ed.). Smithson. Contrib.
Paleobiol. 27.

CLARK, A. H. 1905a. The West Indian parrots. Auk
22:337?344.

CLARK, A. H. 1905b. The Greater Antillean macaws.
Auk 22:345?348.

COLLAR, N. J. 1997. Family Psittacidae. Pp. 280?
477 in Handbook of the birds of the world, Vol.
4. (del Hoya, J., A. Elliott, and J. Sargatal, eds.).
Barcelona, Spain: Lynx Ediciones.

CROZIAT, L., G. NELSON, AND D. E. ROSEN. 1974.
Centers of origin and related concepts. Syst. Zool.
23:265?287.

CUELLO, J. P. 1988. Lista de las aves fósiles de la
región neotropical y de las islas antillanas. Paula-
Coutiana 2:3?79.

DARLINGTON, P. J., JR. 1957. Zoogeography: the
geographical distribution of animals. New York:
John Wiley & Sons, Inc.

DENNY, W. 1847a. A few remarks on the geographical
distribution of birds in the West Indies.
Proc. Zool. Soc. Lond. 15:36?41.

DENNY, W. 1847b. A few remarks on the geographical
distribution of birds in the West Indies.
Ann. Mag. Nat. Hist. 19(29):464?469.

DESJARDINS, P., AND R. MORAIS. 1990. Sequence
and gene organization of the chicken mitochondrial genome. J. Mol. Biol. 212:599?634.

DÍAZ FRANCO, S. 1999. Dos registros nuevos de
aves endémicas en depósitos fosilíferos de Cuba.
Pitirre 12:12?13.

DOMNING, D. P., R. J. EMRY, R. W. PORTELL, S. K.
DONOVAN, AND K. S. SCHINDLER. 1997. Oldest
West Indian land mammal: rhinocerotoid ungulate
from the Eocene of Jamaica. J. Vert. Paleontol.
17:638?641.

DONNELLY, T. W. 1988. Geologic constraints on
Caribbean biogeography. Pp. 15?37 in Zoogeography
of Caribbean insects (Liebherr, J. K.,
ed.). Ithaca, NY: Cornell Univ. Press.

DU TERTRE, LE R. P. J.-B. 1654. Histoire générale
des iles de S. Christophe, de la Guadeloupe, de la
Martinique, et autres dans l?Amérique. Paris: J.
Langlois et E. Langlois.

DU TERTRE, LE R. P. J.-B. 1667. Histoire générale
des Antilles habitées par les Français. Paris: T.
Iolly.

EDWARDS, S. V., AND A. C. WILSON. 1990. Phylogenetically
informative length polymorphism
and sequence variability in mitochondrial DNA
of Australian songbirds (Pomatostomus). Genetics
126:695?711.

EDWARDS, S. V., P. ARCTANDER, AND A. C. WILSON.
1991. Mitochondrial resolution of a deep
branch in the genealogical tree for perching birds.
Proc. Zool. Soc. Lond., Ser. B 243:99?107.

EMSLIE, S. D., AND G. S. MORGAN. 1994. A catastrophic
death assemblage and paleoclimatic implications
of Pliocene seabirds of Florida. Science
264:684?685.

ENKERLIN-HOEFLICH, E. C. 1995. Comparative
ecology and reproductive biology of three species
of Amazona parrots in northeastern Mexico. Ph.
D. diss., Texas A&M University.

FEDUCCIA, A. 1995. Explosive evolution in the Tertiary
birds and mammals. Science 267:637?638.

FEDUCCIA, A. 1996. The origin and evolution of
birds. New Haven, CT: Yale Univ. Press.

FELSENSTEIN, J. 1985. Confidence limits on phylogenies:
an approach using the bootstrap. Evolution
39:783?791.

FORSHAW, J. M. 1989. Parrots of the world, 3rd ed.
Melbourne, Australia: Landsdowne Eds.

GNAM, R. S. 1990. Zur Biologie der Bahama ?
Amazone Amazona leucocephala bahamensis auf
Great Inagua. Papageien 2:89-92.

GNAM, R. S. 1991. Breeding biology of the Bahama Parrot (Amazona leucocephala bahamensis). Ph.
D. diss., City Univ. New York.

GNAM, R., AND A. BURCHSTED. 1991. Population
estimates for the Bahama Parrot on Abaco Island,
Bahamas. J. Field Ornithol. 62:139?146.

GNAM, R., AND R. F. ROCKWELL. 1991. Reproductive
potential and output of the Bahama Parrot,
Amazona leucocephala bahamensis. Ibis
133:400?405.

GNAM, R. S., M. WILSON, AND I. LOTHIAN. 1995.
Breeding biology of the Bahama parrot on Great
Inagua. Pitirre 10(1):26.

GOTELLI, N. J., AND G. R. GRAVES. 1990. Body
size and the occurrence of avian species on landbridge
islands. J. Biogeogr. 17:315?325.

GRAYBEAL, A. 1998. Is it better to add taxa or characters
to a difficult phylogenetic problem? Syst.
Biol. 1:9?17.

HAFFER, J. 1985. Avian zoogeography in the
neotropical lowlands. in Neotropical ornithology
(Buckley, P. A., M. S. Foster, E. S. Morton, R. S.
Ridgely, and F. G. Buckley, eds.). American Ornithologists
? Union. Ornithol. Monogr. 36:113?
146.

HAQ, B. V., J. HARDENBOL, AND P. R. VAIL. 1987.
Chronology of fluctuating sea levels since the
Triassic. Science 235:1156?1167.

HASEGAWA, M., AND H. KISHINO. 1994. Accuracies
of the simple methods for estimating the bootstrap
probability of a maximum-likelihood tree.
Mol. Biol. Evol. 11:142?145.

HEARTY, P. J., P. KINDLER, H. CHENG, AND R. L.
EDWARDS. 1999. A +20 m middle Pleistocene
sea-level highstand (Bermuda and the Bahamas)
due to partial collapse of Antarctic ice. Geology
27:375?378.

HEDGES, S. B. 1996. Historical biogeography of
West Indian vertebrates. Annu. Rev. Ecol. Syst.
27:163?196.

HEDGES, S. B., C. A HASS, AND L. R. MAXSON.
1992. Caribbean biogeography: molecular evidence
for dispersal in West Indian terrestrial vertebrates.
Proc. Natl. Acad. Sci. USA 89:1909?
1913.

HELM-BYCHOWSKI, K., AND J. CRACRAFT. 1993.
Recovering phylogenetic signal from DNA sequences:
relationships within the Corvine assemblage
(Class Aves) as inferred from complete sequences
of the mitochondrial DNA cytochrome-b
gene. Mol. Biol. Evol. 10:1196?1214.

HILLIS, D. M., AND J. J. BULL. 1993. An empirical
test of bootstrapping as a method for assessing
confidence in phylogenetic analysis. Syst. Biol.
42:182?192.

HILLIS, D. M. 1998. Taxonomic sampling, phylogenetic
accuracy, and investigator bias. Syst. Biol.
47:3?8.

HUNT, J. S., E. BERMINGHAM, AND R. E. RICKLEFS.
2001. Molecular systematics and biogeography of
Antillean thrashers, tremblers, and mockingbirds
(Aves: Mimidae). Auk 118:35?55.

IRWIN, D. M., T. D. KOCHER, AND A. C. WILSON.
1991. Evolution of the cytochrome b gene in
mammals. J. Mol. Evol. 32:128?144.

ITURRALDE-VINENT, M. A., AND R. D. E.
MACPHEE. 1999. Paleogeography of the Caribbean
region: implications for Cenozoic biogeography.
Bull. Am. Mus. Nat. Hist. 238:1?95.

JAMES, H. F. 1995. Prehistoric extinctions and ecological
changes on oceanic islands. Ecolog. Stud.
115: 87?102.

JUNIPER, T., AND M. PARR. 1998. Parrots: a guide
to parrots of the world. New Haven, CT: Yale
Univ. Press.

KISHINO, H., AND M. HAGESAWA. 1989. Evaluation
of the maximum likelihood estimate of the evolutionary
tree topologies from DNA sequence data,
and the branching order in Hominoidea. J. Mol.
Evol. 29:170?179.

KLEIN, N. K., AND W. M. BROWN. 1994. Intraspecific
molecular phylogeny in the Yellow Warbler
(Dendroica petechia), and implications for avian
biogeography in the West Indies. Evolution
48:1914?1932.

KLICKA, J., AND R. M. ZINK. 1997. The importance
of recent Ice Ages in speciation: a failed paradigm.
Science 277:1666?1669.

KLUGE, A. G. 1988. Parsimony in vicariance biogeography:
a quantitative method and a Greater
Antillean example. Syst. Zool. 37:315?328.

KOCHER, T. D., W. K. THOMAS, A. MEYER, S. V.
EDWARDS, S. PÄÄBO, F. X. VILLABLANCA, AND
A. C. WILSON. 1989. Dynamics of mitochondrial
DNA evolution in animals: amplification and sequencing
with conserved primers. Proc. Natl.
Acad. Sci. USA 86:6196?6200.

KOENIG, S. 1999. The reproductive biology of Jamaica
?s Black-billed Parrot (Amazona agilis) and
conservation implications. Ph.D. diss., Yale
Univ., New Haven, CT.

KOENIG, S. 2001. The breeding biology of Blackbilled
Parrot Amazona agilis and Yellow-billed
Parrot Amazona collaria in Cockpit Country, Jamaica.
Bird Conserv. Intern. 11:205?225.

LABAT, J. B. 1722. Nouveau voyage aux isles de
l?Amérique. Paris: La Haye, P. Husson.

LABAT, J. B. 1724. Nouveau voyage aux isles de
l?Amérique. Vols. I & II. Paris: La Haye, P. Husson,
T. Johnson, P. Gosse, J. van Duren, R. Alberts,
C. Le Vier.

LABAT, J. B. 1742. Nouveau voyage aux isles de
l?Amérique. New ed. Paris: La Haye, P. Husson.

LACK, D. 1976. Island biology, illustrated by the
land birds of Jamaica. Berkeley: Univ. Calif.
Press.

LAKE, J. A. 1994. Reconstructing evolutionary trees
from DNA and protein sequences: paralinear distances.
Proc. Natl. Acad. Sci. USA 9:1455?1459.

LANTERMANN, W. 1997. Verbreitung und evolution
der psittacidenfauna auf den ozeanischen inseln
der Karibischen See. Papageienkunde 1:263?278.

LEETON, P., L. CHRISTIDIS, AND M. WESTERMAN.
1993. Feathers from museum bird skins ? a
good source of DNA for phylogenetic studies.
Condor 95:465?466.

LÉOTAUD, A. 1866. Oiseaux de l?île de La Trinidad,
(Antilles). Port-d?Espagne: Chronicle Publ. Off.

LITTELL, R. 1993. Controlled wildlife: Federal permit
procedures, Vol. 1, 2nd ed. Washington DC:
Assoc. Syst. Collections.

LOCKHART, P. J., M. A. STEEL, M. D. HENDY, AND
D. PENNY. 1994. Recovering evolutionary trees
under a more realistic model of sequence evolution.
Mol. Biol. Evol. 11:605?612.

LOUSADA, S. A., AND S. N. G. HOWELL. 1996. Distribution,
variation, and conservation of Yellowheaded
Parrots in northern Central America.
Cotinga 5:46?53.
MADDISON, W. P., AND D. R. MADDISON. 1992.
MacClade: analysis of phylogeny and character
evolution. Version 3. Sunderland, MA: Sinauer.

MALFAIT, B., AND M. DINKELMANN. 1972. Circum-
Caribbean tectonics and igneous activity and the
evolution of the Caribbean plate. Geol. Soc. Am.
Bull. 83:251?272.

MARIEN, D., AND K. F. KOOPMAN. 1955. The relationships
of West Indian species of Aratinga
(Aves, Psittacidae). Am. Mus. Nat. Hist. Novit.
1712:1?20.

MEDLIN, L., H. J. ELWOOD, S. STICKEL, AND M. L. SOGIN. 1988. The characterization of enzymatically
amplified eukaryotic 16S-like rRNA coding
regions. Gene 71:491?499.

MINDELL, D. P., AND C. E. THACKER. 1996. Rates
of molecular evolution: phylogenetic issues and
applications. Annu. Rev. Ecol. Syst. 27:279?303.

MIYAKI, C. Y., S. R. MATIOLI, T. BURKE, AND A.
WAJNTAL. 1998. Parrot evolution and paleogeographical
events: mitochondrial DNA evidence.
Mol. Biol. Evol. 15:544?551.

MORGAN, G. S. 1989. Fossil Chiroptera and Rodentia
from the Bahamas, and the historical biogeography
of the Bahamian mammal fauna. Pp. 685?
740 in Biogeography of the West Indies: past,
present, and future (Woods, C. A., ed.). Gainesville,
FL: Sandhill Crane Press.

MORGAN, G. S. 1994. Late Quaternary fossil vertebrates
from the Cayman Islands. Pp. 465?508 in
The Cayman Islands: natural history and biogeography
(Brunt, M. A., and J. E. Davies, eds.).
Dordrecht, Netherlands: Kluwer Academic Publishers.

MORGAN, G. S. 2001. Patterns of extinction in West
Indian bats. Pp. 369?407 in Woods, C. A., and F.
E. Sergile, eds. Biogeography of the West Indies:
patterns and perspectives, 2nd ed. Boca Raton, FL:
CRC Press.

MORGAN, G. S., AND C. A. WOODS. 1986. Extinction
and the zoogeography of West Indian land
mammals. Biol. J. Linn. Soc. 28:167?203.

OLSON, S. L. 1977. Pleistocene birds of Puerto
Rico. Natl. Geogr. Res. Rep. 18:563?566.

OLSON, S. L. 1978. A paleontological perspective
of West Indian birds and mammals. Pp. 99?117
in Zoogeography in the Caribbean (Gill, F. B.,
ed.). Spec. Publ. 13. Philadelphia, PA: Acad. Nat.
Sci.

OLSON, S. L. 1982. Biological archeology in the
West Indies. Fla. Anthropol. 35:162?168.

OLSON, S. L. 1985. The fossil record of birds. Pp.
79?239 in Avian biology, Vol. VIII (Farner, D.
S., J. R. King, and K. C. Parkes, eds.). Orlando,
FL: Academic Press, Inc.

OLSON, S. L. 1989. Aspects of global avifaunal dynamics
during the Cenozoic. Pp. 2023?2029 in
Symposium 35: The early radiation of birds
(Kurochkin, E. N., and S. L. Olson, eds.). Acta
XIX Congressus Internationalis Ornithologici 2.
Ottawa, Canada.

OLSON, S. L., AND W. B. HILGARTNER. 1982. Fossil
and subfossil birds from the Bahamas. Pp. 22?56 in Fossil Vertebrates from the Bahamas
(Olson, S. L., ed.). Smithson. Contrib. Paleobiol.
48.

OLSON, S. L., AND G. K. PREGILL. 1982. Introduction
to the paleontology of Bahamian vertebrates.
Pp. 1?7 in Olson, S. L., ed. Fossil vertebrates
from the Bahamas: Smithson. Contrib. Paleobio.
48.

PAYNTER, R. A., JR. 1955. The ornithogeography of
the Yucatán Peninsula. Bull. Peabody Mus. Nat.
Hist. 9:1?347.

PINDELL, J. L., S. C. CANDE, W. C. PITMAN, III, D.
B. ROWLEY, J. F. DEWEY, J. LABRECQUE, AND
W. HAXBY. 1988. A plate-kinematic framework
for models of Caribbean evolution. Tectonophysics
155:121?138.

PORTELL, R. W., S. K. DONOVAN, AND D. P.
DOMNING. 2001. Early Tertiary vertebrate fossils
from Seven Rivers, Parish of St. James, Jamaica,
and their biogeographical implications. Pp. 191-
200 in Biogeography of the West Indies: patterns
and perspectives, 2nd ed. (Woods, C. A., and F. E.
Sergile, eds.). Boca Raton, FL: CRC Press.

POSADA, D., AND K. A. CRANDALL. 1998.
MODELTEST: testing the model of DNA substitution.
Bioinformatics 14:817?818.

PREGILL, G. K. 1981. An appraisal of the vicariance
hypothesis of Caribbean biogeography and its
application to West Indian terrestrial vertebrates.
Syst. Zool. 30:147?155.

PREGILL, G. K., AND S. L. OLSON. 1981. Zoogeography
of West Indian vertebrates in relation
to Pleistocene climatic cycles. Annu. Rev. Ecol.
Syst. 12:75?98.

PREGILL, G. K., D. W. STEADMAN, S. L. OLSON,
AND F. V. GRADY. 1988. Late Holocene fossil
vertebrates from Burma Quarry, Antigua, Lesser
Antilles. Smithson. Contrib. Zool. No. 463.

PREGILL, G. K., D. W. STEADMAN, AND D. R.
WATTERS. 1994. Late Quaternary vertebrate faunas
of the Lesser Antilles: historical components
of Caribbean biogeography. Bull. Carnegie Mus.
Nat. Hist. 30:1?51.

RAFFAELE, H., J. WILEY, O. GARRIDO, A. KEITH,
AND J. RAFFAELE. 1998. A guide to the birds of
the West Indies. Princeton, NJ: Princeton Univ.
Press.

RICKLEFS, R. E., AND G. W. COX. 1972. Taxon cycles
in the West Indian avifauna. Am. Nat. 106
(948):195?219.

RICKLEFS, R. E., AND G. W. COX. 1978. Stage of taxon cycle, habitat distribution, and population
density in the avifauna of the West Indies. Am.
Nat. 112(987):875?895.

ROSEN, D. E. 1976. A vicariance model of Caribbean
biogeography. Syst. Zool. 24:431?464.

SANGER, F., S. NICKLEN, AND A. R. COULSON.
1977. DNA sequencing with chain-terminating
inhibitors. Proc. Natl. Acad. Sci. USA 74:5463?
5467.

SCHUCHMANN, K.?L. 1980. Ökologie und Evolution
der Trochilidenfauna auf den ozeanischen
Inseln der Karibischen See. Bonn. Zool. Beitr.
31:289?309.

SCLATER, P. L. 1891. On recent advances in our
knowledge of the geographical distribution of
birds. Ibis 3(12):514?557.

SCHRÖDER, V. W. 1988. Zur Biologie und zum
status der Kubaamazone (Amazona leucocephala)
auf Great Abaco (Bahamas). Trochilus 9:3?34.

SEUTIN, G., N. KLEIN, R. E. RICKLEFS, AND E. BERMINGHAM.
1994. Historical biogeography of the
Bananaquit (Coereba flaveola) in the Caribbean
region: a mitochondrial DNA assessment. Evolution
48:1041-1061.

SHIELDS, G. F. AND A. C. WILSON. 1987. Calibration
of mitochondrial DNA evolution in geese. J.
Mol. Evol. 24:212-217.

SLIKAS, B., S. L. OLSON, AND R. C. FLEISCHER.
2002. Rapid independent evolution of flightlessness
in four species of Pacific island rails
(Rallidae), an anlysis based on mitochondrial sequence
data. J. Avian Biol. 33:5?14.

SMITH, G.A. 1975. Systematics of parrots. Ibis
117:18?68.

SNYDER, N. F. R., W. B. KING, AND C. B. KEPLER.
1982. Biology and conservation of the Bahama
Parrot. Living Bird 19:91?114.

SNYDER, N. F. R., J. W. WILEY, AND C. B. KEPLER.
1987. The parrots of Luquillo: natural history and
conservation of the Puerto Rican Parrot. Los Angeles,
CA: West. Found. Vert. Zool.

SORENSON, M. D., AND T. W. QUINN. 1998. Numts:
a challenge for avian systematics and population
biology. Auk 115:214?221.

STANGEL, P. W., AND M. R. LENNARTZ. 1988. Survival
of Red-cockaded Woodpecker nestlings unaffected
by sampling blood and feather pulp for
genetic studies. J. Field Ornithol. 59:389?394.

STEADMAN, D. W., AND G. S. MORGAN. 1985. A
new species of bullfinch (Aves: Emberizinae) from a late Quaternary cave deposit on Cayman
Brac, West Indies. Proc. Biol. Soc. Wash.
98:544?553.

STEADMAN, D. W., G. K. PREGILL, AND S. L. OLSON.
1984. Fossil vertebrates from Antigua,
Lesser Antilles: evidence for late Holocene human-
caused extinctions in the West Indies. Proc.
Natl. Acad. Sci. USA 81:4448?4451.

SWOFFORD, D. L. 1998. PAUP*: Phylogenetic
analysis using parsimony (*and other methods),
Version 4. Sunderland, MA: Sinauer Associates.

SWOFFORD, D. L., G. J. OLSEN, P. J. WADDELL,
AND D. M. HILLIS. 1996. Phylogenetic inference.
Pp. 407?514 in Molecular systemtatics (Hilllis,
D. M., C. Mortiz, and B. K. Mable, eds.). Sunderland,
MA: Sinauer Associates, Inc.

TARR, C. L. AND R. C. FLEISCHER. 1993. Mitochondrial-
DNA variation and evolutionary relationships
in the Amakihi complex. Auk 110:825-831.

TERBORGH, J., J. FAABORG, AND H. J. BROCKMANN.
1978. Island colonization by Lesser Antillean
birds. Auk 95:59?72.

THOMPSON, D. W. 1899. On characteristic points in
the cranial osteology of the parrots. Proc. Zool.
Soc. Lond. 1899:9?46.

THOMPSON, J. D., D. G. HIGGINS, AND T. J. GIBSON.
1994. Clustal W: improving the sensitivity
of progressive multiple sequence alignment
through sequence weighting, position specific gap
penalties, and weight matrix choice. Nucleic Acids
Res. 22:4673?4680.

VALENTINE, M. 1990. Chromosome analysis. Pp.
127?131 in Genus Amazona (Stoodley, J., and P.
Stoodley, eds.). Portsmouth, England: Bezels
Publ.

WEBB, T., III, AND P. J. BARTLEIN. 1992. Global
changes during the last 3 millions years: climatic
controls and biotic responses. Annu. Rev. Ecol.
Syst. 23:141?173.

WEGE, D. C., AND N. J. COLLAR. 1991. The Bluecheeked
Amazon Amazona dufresniana: a review.
Bird Conserv. Intern. 1:317?328.

WETMORE, A. 1917. The birds of Culebra Island,
Porto Rico. Auk 34:51?62.

WETMORE, A. 1926. Descriptions of additional fossil birds from the Miocene of Nebraska. Am.
Mus. Nat. His. Novit. 211:1?5.

WETMORE, A. 1928. Bones of birds from the Ciego
Montero deposit of Cuba. Am. Mus. Nat. His.
Novit. 301:1?5.

WETMORE, A. 1937. Ancient records of birds from
the island of St. Croix with observations on extinct
and living of Puerto Rico. J. Agric. Univ.
Puerto Rico 21:5?16.

WETMORE, A. 1938. Bird remains from the West
Indies. Auk 55:51?55.

WETMORE, A. 1951. Recent additions to our knowledge
of prehistoric birds 1933?1949. Pp. 51?74
in Proc. Tenth Intern. Ornithol. Congr. (1950)
(Hörstadins, S., ed.). Uppsala, Sweden.

WETMORE, A. 1956. A check-list of the fossil and
prehistoric birds of North America and the West
Indies. Smithson. Misc. Coll. 131(5):1?105.

WILEY, J. W. 1991. Status and conservation of parrots
and parakeets in the Greater Antilles, Bahama
Islands, and Cayman Islands. Bird Conserv.
Intern. 1:187?214.

WILEY, J. W. 2000. A bibliography of ornithhology
in the West Indies. Proc. West. Found. Vert.
Zool. 7:1?817.

WILLIAMS, M. I. 1998. The systematic relationships
of the Hawk-headed Parrot, Deroptyus accipitrinus.
in Abstracts of the 1998 North American
Ornithological Conference. St. Louis, MO.

WILLIAMS W., AND D. W. STEADMAN. 2001. The
historic and prehistoric distribution of parrots
(Psittacidae) in the West Indies. Pp. 175?189 in
Biogeography of the West Indies: patterns and
perspectives, 2nd ed. (Woods, C. A., and F. E.
Sergile, eds.). Boca Raton, FL: CRC Press.

WILSON, A. C., R. L. CANN, S. M. CARR, M.
GEORGE, U. B. GYLLENSTEN, K. M. HELMBYCHOWSKI,
R. G. HIGUCHI, S. R. PALUMBI, E.
M. PRAGER, R. D. SAGE, AND M. STONEKING.
1985. Mitochondrial DNA and two perspectives
on evolutionary genetics. Biol. J. Linn. Soc.
26:375?400.

ZUCKERKANDL, E., AND L. PAULING. 1965. Molecules
as documents of evolutionary history. J.
Theor. Biol. 8:357?366.