ORNITOLOGIA NEOTROPICAL ______Volume 16 2005 No. 4 ______

ORNITOLOGIA NEOTROPICAL 16: 449–470, 2005 © The Neotropical Ornithological Society

THE ICTERUS IN THE WEST INDIES

Orlando H. Garrido1, James W. Wiley2,4 , & Arturo Kirkconnell3

1Calle 60 & 1706 entre 17 y 19, Marianao 13 (Playa), La Habana, Cuba. 2Maryland Cooperative Fish & Wildlife Research Unit, University of Maryland Eastern Shore, 1120 Trigg Hall, Princess Anne, Maryland 21853, USA. E-mail: [email protected] 3Museo Nacional de Historia Natural, Calle Obispo No. 61, Plaza las Armas, Ciudad La Habana, Cuba.

Resumen. – El género Icterus en las Antillas. – Examinamos especímenes de las especies endémicas del genero Icterus en las Antillas, así como de los I. prosthemelas y I. prosthemelas praecox, junto con la información disponible sobre su historia natural, suplida con nuestras observaciones de campo en las islas. Determina- mos que la morfología, historia natural, vocalización, y las diferencias moleculares, así como el patrón de distribución entre las islas, definen claramente las formas de Icterus de las Antillas como miembros de una superspecie, con alloespecies distintas en las diversas islas. Por lo tanto, proponemos que éstas alloespecies de las Antillas constituyen especies endémicas de las islas respectivas, y así proponemos el arreglo siguiente: I. northropi (Andros y Abaco), I. melanopsis (Cuba, Isla de Pinos y algunos cayos de la zona norte), I. dominicensis (La Española, incluyendo Île de la Gonâve, Île de la Tortue, Île-à-Vache, y Isla Saona), I. porto- ricensis (Puerto Rico), I. oberi (Montserrat), I. bonana (Martinique), y I. laudabilis (St. Lucia). Abstract. – We examined specimens of endemic Icterus in the West Indies, as well as I. prosthemelas and I. prosthemelas praecox, along with available information on their natural history, supplemented with our field observations in the islands. We determined that morphological, natural history, vocalization, and molecular differences, as well as the pattern of distribution among islands, clearly define the West Indian Icterus forms as members of a superspecies, with distinct allospecies in the different islands. Therefore, we propose that these West Indian allospecies constitute endemic species from their respective islands, and propose the following arrangement: Icterus northropi (Andros and Abaco), I. melanopsis (Cuba, Isla de Pinos, and some northern keys), I. dominicensis (Hispaniola, including Île de la Gonâve, Île de la Tortue, Île-à- Vache, and Isla Saona), I. portoricensis (Puerto Rico), I. oberi (Montserrat), I. bonana (Martinique), and I. lau- dabilis (St. Lucia). Accepted 25 May 2005. Key words: Greater Antillean Oriole, Icterus dominicensis, Icterus melanopsis, Icterus northropi, Icterus portoricensis, , West Indies. ______4Corresponding author.

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INTRODUCTION be conspecific. In his biogeographic analysis of West Indian , Lack (1976) suggested The family Icteridae is represented in the ancestral I. dominicensis from Central America West Indies by nine genera: Dolichonyx, Age- colonized the Greater Antilles, then spread laius, Nesopsar, Sturnella, Xanthocephalus, Dives, into the Lesser Antilles, subsequently evolv- Quiscalus, Molothrus, and Icterus. Of these, the ing into the three present-day species. Most best represented is Icterus, with 12 species and recently, Lovette et al. (1999) examined the subspecies. Only three of these forms are not Lesser Antillean oriole species, using mito- resident breeders: I. spurius, transient in Cuba, chondrial DNA sequences to determine rela- Jamaica, and some of the Bahama Islands, and tionships among these populations, and vagrant in Hispaniola; I. cucullatus, casual or concluded that I. oberi, I. bonana, and I. laudabi- vagrant in Cuba; and I. galbula, transient and lis are evolutionarily significant units that winter resident in the Greater Antilles, and meet species criteria under the phylogenetic vagrant in some Lesser Antilles. West Indian species concept. Because of the consistently Icterus species and subspecies are resident as large mitochondrial divergences among follows: I. dominicensis northropi Allen in The Lesser Antillean Icterus populations, Lovette et Bahamas; I. d. melanopsis (Wagler) in Cuba, Isla al. (1999) concluded that the Lesser Antillean de Pinos (now Isla de la Juventud), and some oriole populations are as genetically distinct as northern keys; I. d. dominicensis Linnaeus in many avian species. Hispaniola; I. d. portoricensis Bryant in Puerto Until recently, I. dominicensis consisted of Rico; I. laudabilis Sclater in St. Lucia; I. oberi populations from southern Veracruz, north- Lawrence in Montserrat; I. bonana (Linnaeus) ern Oaxaca, Tabasco, Chiapas, and the in Martinique; I. icterus (Linnaeus), introduced Yucatan Peninsula south on the and established in Puerto Rico, Mona, St. slope of Central America to extreme western Thomas, Jamaica, St. John, Antigua, Domin- Panama, and The Bahamas and West Indies ica, Grenada, and Trinidad; and I. leucopteryx (A. O. U. 1998). Using a molecular analysis of (Wagler) in Jamaica, Grand Cayman, and San phylogeny, Omland et al. (1999) reached the Andrés. Currently, only I. bonana, I. oberi, and conclusion that the forms of Icterus pros- I. laudabilis are considered endemic species (A. themelas (Black-cowled Oriole) and the domi- O. U. 1998, Raffaele et al. 1998). With the nicensis group (Greater Antillean Oriole; exception of I. leucopteryx and I. icterus, there is including dominicensis, melanopsis, portoricensis, general agreement that all resident West and northropi) were distinct species, which was Indian Icterus are within the I. dominicensis subsequently accepted by the A. O. U. (2000: superspecies complex (Lovette et al. 1999). 853). Omland et al. (1999) had the impression The taxonomic status of the various Antillean that some of the subspecies included in the island populations, however, has been long dominicensis group also could be distinct spe- disputed (Blake 1968, Sibley & Monroe 1990). cies, rather than subspecies, but they did not In support of his hypothesis that the have suitable information on natural history, Lesser Antillean species evolved from Greater vocal behavior, and plumage differences to Antillean ancestors, Bond (1956) considered elaborate on those relationships. Therefore, the Lesser Antillean I. oberi, I. bonana, and I. the four West Indian taxa were retained as laudabilis as derived “representatives of I. conspecific (A. O. U. 2000). dominicensis.” Blake (1968) also considered the The recognition of I. prosthemelas is not Lesser Antillean orioles as derived from I. new. Although originally described as a spe- dominicensis, with which he thought they may cies by Strickland (1850) under the genus

450 ICTERUS IN THE WEST INDIES

Xanthornus, Hellmayr (1937) considered it We present summary descriptive statistics conspecific with the Bahaman population (mean, SD, and range) for the specimens. northropi Allen, 1890, as I. prosthemelas northropi, Unpaired t-tests were used to test for equality and separate from the subspecies dominicensis, of means among populations. We used Bon- melanopsis, and portoricensis. Conversely, Bond ferroni’s correction in conducting multiple (1938) considered northropi distinct from pros- tests of means. Further, we used ANOVA themelas and derived from Cuban melanopsis. analyses to determine if differences exist Phillips & Dickerman (1965) described the among island populations. Body measure- species praecox from Panamá and Costa Rica, ments were plotted to assess the pattern of recognizing I. prostemelas as a distinct species. spatial segregation among populations. The Blake (1968), however, retained the two Cen- hypothesis of separation derived from the tral American populations as subspecies, I. d.