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Tyrannus Caudifasciatus) Complex (Aves: Tyrannidae)

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Tyrannus Caudifasciatus) Complex (Aves: Tyrannidae) The Wilson Journal of Ornithology 121(4):703–713, 2009 TAXONOMY OF THE LOGGERHEAD KINGBIRD (TYRANNUS CAUDIFASCIATUS) COMPLEX (AVES: TYRANNIDAE) ORLANDO H. GARRIDO,1 JAMES W. WILEY,2,4 AND GEORGE B. REYNARD3 ABSTRACT.—We examined the complex of populations of the Loggerhead Kingbird (Tyrannus caudifasciatus), a West Indian endemic. We separate populations in Puerto Rico and Isla Vieques (T. taylori), and Hispaniola (T. gabbii) as distinct species. Subspecific distinction is assigned to populations in Cuba, Isla de Pinos, and Cuban satellites (T. caudifasciatus caudifasciatus); Cayman Islands (T. c. caymanensis); Jamaica (T. c. jamaicensis); and the Bahamas (T. c. bahamensis)on the basis of differences in plumage coloration and pattern, size, vocalizations, and distribution. Received 1 December 2008. Accepted 27 June 2009. The Loggerhead Kingbird (Tyrannus caudifas- man; P. taylori from Puerto Rico, P. bahamensis ciatus) is endemic to the Bahama Islands and from the Bahamas, and P. gabbii from Hispaniola. Greater Antilles, although occasionally stragglers Sharpe (1901) separated T. jamaicensis of Ja- have been reported in southern Florida (Bond maica. Ridgway (1907:677–678) noted differenc- 1978:3, Robertson and Woolfenden 1992, AOU es among populations of Loggerhead Kingbird 1998: 414). Tyrannus caudifasciatus was origi- based on several characteristics, including the nally described from Cuba by d’Orbigny in 1839 extent to which the bases of the toes are united: in in the genus Tyrannus. Subsequent authors used T. bahamensis and T. caudifasciatus half of basal the genus Pitangus and, later, Tolmarchus Ridg- phalanx of the middle toe is united to the outer way (1905) with T. taylori of Puerto Rico as the toe, whereas in T. taylori more than half of the type species. The morphological differences that basal phalanx is united to the outer toe. Cory and separate Tolmarchus from Tyrannus are slight. Hellmayr (1927) and Bond (1936, 1956) consid- Both groups are generally similar in voice and ered Tyrannus caudifasciatus to be a polytypic nesting behavior and, according to Bond species with the subspecies T. c. caymanensis (1956:105), in not having the outer primaries (Nicoll 1904) designated for the Cayman Islands, attenuated at their tips. Meise (1949), Bond T. c. jamaicensis (Chapman 1892) for Jamaica, T. (1958), and later authors merged Tolmarchus with c. taylori (Sclater 1864 in Taylor 1864) for Puerto Tyrannus. Smith (1966) presented characteristics Rico and Isla Vieques, T. c. bahamensis (Bryant of vocalizations of some species of Tyrannus, 1864) for some of the northern Bahama Islands, T. including the Loggerhead Kingbird, suggesting c. caudifasciatus for Cuba and Isla de Pinos, and there were four species groups within the genus. T. c. gabbii (Lawrence 1876) for Hispaniola. Some workers, however, maintain T. caudifascia- Wetmore and Swales (1931:305), however, wrote, tus in a monotypic subgenus Tolmarchus (Bond ‘‘We do not agree with [Cory and] Hellmayr 1967). Lack (1976: 297), comparing T. caudifas- (1927) in considering gabbii a subspecies of ciatus with the other two Greater Antillean Tolmarchus caudifasciatus as its characters seem Tyrannus species (Giant Kingbird [T. cubensis] to entitle it to specific recognition.’’ Parkes and Grey Kingbird [T. dominicensis]), noted, ‘‘the (1963) separated the Isla de Pinos population relationships of these three species to each other from T. c. caudifasciatus under the subspecies T. and to T. tyrannus are not clear.’’ c. flavescens, an arrangement followed by Traylor Cory (1889, 1892) considered the then-known (1979). Dickinson (2003) maintained the subspe- forms of the Loggerhead Kingbird to be separable cific level for all populations of T. caudifasciatus, at the species level, recognizing Pitangus caudi- including T. c. flavescens of Isla de Pinos. fasciatus from Cuba, Jamaica, and Grand Cay- Little has been written concerning the taxono- my of the Tyrannus caudifasciatus complex with the exception of Ridgway (1907), Wetmore and 1 Calle 60 Nu´mero 1706, entre 17 y 19, Marianao 13 Swales (1931), and Parkes (1963). We present the (Playa), La Habana, Cuba. results of our comparison of morphology and 2 Western Foundation of Vertebrate Zoology, P. O. Box 64, Marion Station, MD 21838, USA. vocalizations among populations in an effort to 3 Deceased. establish species limits within the Loggerhead 4 Corresponding author; e-mail: [email protected] Kingbird. 703 704 THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 121, No. 4, December 2009 METHODS webs of rectrices passing gradually into dull We measured 711 Loggerhead Kingbird spec- whitish at extreme base; tail not tipped with imens in 11 United States and Cuban institutions. brownish gray or whitish.’’ Ridgway further split Specimens from the Cuban cays included cayos this second group, using the following characters: Cinco Leguas, France´s, Las Brujas, Santa Marı´a, ‘‘Smaller…; rectrices edged conspicuously with Coco, Paredo´n Grande, Guajaba, and Sabinal in rusty’’ (5 T. gabbii); and ‘‘Larger…; rectrices not the Archipie´lago Sabana–Camagu¨ey. All speci- edged with rusty’’ (5 T. taylori). Another mens were measured with a ruler and dial calipers important character, overlooked by Ridgway, is to the nearest 0.1 mm following Baldwin et al. the shape of the tail, which is obviously notched at (1931). Only adult specimens (n 5 674) were the end (although not as much as in the Grey used in comparing morphometrics within and Kingbird, T. dominicensis)intaylori, and less so among populations. Chord measurements were in bahamensis, whereas the rest of the taxa have made with the wing flattened against the ruler. square-tipped tails. Culmen measurements are of the exposed portion We present a comparative diagnosis of the (without cere). Most mass data were obtained plumage of the taxa, as well as a synopsis of some from post-mortem birds. We include mass data of of these characters (Table 1) based on examina- Steadman et al. (1980) and Olson et al. (1981) in tion of 711 specimens. addition to our mass data. Tyrannus caudifasciatus bahamensis (Baha- We present summary descriptive statistics mas).—Head with a browner tinge and not as (mean, SD, and range) for the specimens. black as in other forms, clearly contrasting with the grayish upperparts, which show a brownish Unpaired t-tests were used to test for equality of suffusion. Concealed crown patch generally a mix weights among populations. We used Bonferro- of orange and yellow. Underparts, including ni’s correction in conducting multiple tests of axillae, with considerable yellow (more than in means and ANOVA analyses to test differences any other population). No brown suffusion on among island populations. Body measurements breast. Edge of wing coverts grayish. The blotches were plotted to assess the pattern of spatial of the inner rectrices yellowish, and more segregation among populations. The hypothesis restricted than in caudifasciatus, jamaicensis,or of separation derived from the plots of body caymanensis. Bahamian birds have a rusty rump, a measurements was tested using discriminate character not found in other populations of function analysis (DFA) (Kleinbaum and Kupper caudifasciatus. Tail slightly notched, although 1978). Statistical computations were performed not as much as in taylori, and tipped with buff, using the software package Minitab (2000) not white or gray. including DFA and cluster analyses. We used Tyrannus c. caudifasciatus (Cuba, Isla de Pinos, the single linkage method and squared Euclidean and cays).—Head blackish, strongly contrasting measure to conduct cluster analyses. Level of with the brownish-sepia upperparts. Concealed significance was set at 0.05. crown patch variable yellow, orange, or a mix of Most recordings of vocalizations were obtained both colors; individuals displaying the three color by Reynard and are deposited in his private types may occur in the same population. Edge of collection. Additional vocalization recordings wing coverts whitish. Tail rather squared at tip, were obtained from the Macaulay Library of the tipped in white or light gray, without rust color. Cornell Laboratory of Ornithology and from Inner rectrices with well-developed whitish-yel- Bruce Sorrie. Vocalizations were analyzed using low blotches. No brownish tinge on breast. Raven (Version 1.0) software. Populations in Cuba, Isla de Pinos, and the cays all share these characters. RESULTS Tyrannus caudifasciatus caymanensis (Cayman Plumage Characters.—Ridgway (1907:678) Islands).—Head sooty-slate, rather than black, split the taxa into two groups based on: (Group contrasting with the brownish-gray upperparts, 1) ‘‘Inner webs of rectrices with basal third, or which are more olivaceous than in T. c. caudi- more, abruptly yellowish white; tail tipped with fasciatus. Concealed crown patch between yellow pale brownish gray or whitish’’ (the former and orange in color. More suffusion of yellow on important character was completely overlooked the underparts than in other populations with the by subsequent authors); and (Group 2) ‘‘Inner exception of bahamensis. We noted that birds Garrido et al. N LOGGERHEAD KINGBIRD SPECIES LIMITS 705 TABLE 1. Plumage characteristics of Loggerhead Kingbirds (Tyrannus caudifasciatus) in eight populations. Bold data highlight patterns of similarity. Population Plumage character Bahamas Cuba Isla de Pinos Cuban caysa Jamaica Cayman Islands Hispaniola Puerto Rico Tip of tailb B WW W W W NN Shape of tailc HN SS S S S S N Crown patchd M Y–O Y–O Y–O Y Y–O Y Y Breast featherse AA A A A A P A Inner rectricesf PP P P P P AA Rectricesg NR NR NR NR NR NR R NR Yellow on underpartsh L NN N N MNN a Cuban cays include cayos Cinco Leguas, France´s, Las Brujas, Santa Marı´a, Coco, Paredo´n Grande, Guajaba, and Sabinal in the Archipie´lago Sabana– Camagu¨ey. b Tip of tail: W 5 tipped with whitish, B 5 tipped with buff, N 5 not tipped. c Shape of tail end: N 5 notched, HN 5 half notched, S 5 square. d Crown patch: Y 5 yellow, O 5 orange, M 5 mixed yellow and orange.
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