Geographic Variation
Body size varies geographically: the largest birds breed along the Pacific Coast, birds of intermediate size breed along the Gulf and Atlantic coasts, and the smallest birds breed in the Caribbean (see Appendix 1). Plumage varies less strikingly, but in general breeders in the West Indies are darker than those on the Atlantic and Gulf of Mexico. Soft part coloration (when birds are in breeding condition), particularly the basal portion of the gular, varies red on the Pacific of North America to dark greenish on the Atlantic and in the Caribbean to blackish in w. South America.
Subspecies
Five subspecies, following Wetmore (1945), distinguished on the basis of differences in plumage color, the color of the bill and orbital in breeding condition, and body size. Diagnoses below are adapted from those of Wetmore (1945), although he also relied on hindneck color during the breeding season, a character since shown to vary greatly within and among subspecies (Schreiber et al. 1989). Range limits of subspecies are poorly known, especially in Middle America and n. South America. Angehr and Kushlan (2007) suggested that breeders in Panama may be taxonomically distinct, and Nelson (2006) questioned the validity of the Galapagos subspecies, despite differences in gular coloration.
P. o. occidentalis Linnaeus, 1766. Includes P. fuscus Gmelin, 1789. Breeds in the Caribbean from the s. Bahamas south through the Greater and Lesser Antilles to the coasts of Colombia east to Venezuela and Trinidad (Am. Ornithol. Union 1957, van Halewyn and Norton 1984) [type locality = Jamaica]; wanders to the Gulf coast of Florida, Caribbean coast of Mexico (Wetmore 1945), and along the Atlantic coast of South America to e. Brazil (Patrial 2011). Small (see Appendix 1; male wing < 500 mm, female wing < 485 mm); in breeding plumage, ventrum dark moderately streaked with pale; in breeding condition, gular pouch a blackish blue-green.
P. o. carolinensis Gmelin, 1789. Includes P. albicollis Maynard, 1874. Breeds coastally from Maryland south to s. Florida, in the Gulf of Mexico from s. Florida to s. Texas, in s. Veracruz, at various points around the Yucatan Peninsula (Mexico and Belize), and in Honduras (Am. Ornithol. Union 1957, 1998; Blake 1977); also breeds on Pacific coast of Honduras, Costa Rica, and Panama (Am. Ornithol. Union 1957) [type locality = Charleston, South Carolina]; wanders north along the Atlantic Coast to the Maritime Provinces, south to e. Brazil, inland to various locales in the e. United States and Middle America, and on the Pacific Coast from s. Mexico to n. Peru (Murphy 1936, Am. Ornithol. Union 1957, 1998, D. Blankinship pers. comm). Like P. o. occidentalis, but body size much larger (male wing > 500 mm, female wing > 485 mm) and ventrum of breeding plumage generally paler.
P. o. californicus Ridgway, 1884. Breeds coastally at scattered locations from the Channel Is. off s. California south along the Baja California Peninsula and through the Gulf of California south to Sinaloa (Anderson et al. 2013), and has bred inland at the Salton Sea in se. California (Sturm 1998) [type locality = La Paz, Baja California Sur]; has wandered both coastally north to se. Alaska (Tobish 2003) and south coast to El Salvador and inland to the sw. United States (Am. Ornithol. Union 1957, U.S. Fish Wildl. Serv. 1983), with putative records from the Gulf Coast of Mexico, Texas, and Florida and from the Atlantic Coast of Florida and, possibly, Georgia (Brinkley 2010). Similar to P. o. carolinensis, but gular in breeding condition bright red (Schreiber et al. 1989); averages larger.
P. o. murphyi Wetmore, 1945. Breeds along the Pacific coast and offshore islands of Colombia (Moreno and Buelvas 2005) and Ecuador (Suárez and Calle 2005, U.S. Fish Wildl. Serv. 2009) [type locality = Pelado I., Santa Elena Bay, Ecuador]; ranges along the Pacific coast south to n. Chile (Wetmore 1945, Jaramillo 2007). Like P. o. carolinensis, but dorsum darker and ventrum more extensively streaked with pale (Wetmore 1945).
P. o. urinator Wetmore, 1945. Resident in the Galápagos Is. [type locality = Española (formerly Hood) I.]. Similar to P. o. californicus, but gular blackish (not red; Rothschild and Hartert 1899) and dorsum (and often ventrum, too) darker (Wetmore 1945).