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Short Communications SHORT COMMUNICATIONS The Condor 106:664±670 q The Cooper Ornithological Society 2004 PHYLOGENETIC RELATIONSHIPS OF THE RED-BELLIED GRACKLE (ICTERIDAE: HYPOPYRRHUS PYROHYPOGASTER) INFERRED FROM MITOCHONDRIAL DNA SEQUENCE DATA CARLOS DANIEL CADENA1,4,ANDRE S M. CUERVO2,5 AND SCOTT M. LANYON3 1Department of Biology and International Center for Tropical Ecology, University of Missouri-St. Louis, 8001 Natural Bridge Road, St. Louis, MO 63121 2Instituto de BiologõÂa, Universidad de Antioquia, MedellõÂn, Colombia 3Bell Museum of Natural History and Department of Ecology, Evolution, and Behavior, University of Minnesota, St. Paul, MN 55108 Abstract. The phylogenetic af®nities of the Red- Relaciones FilogeneÂticas de Hypopyrrhus bellied Grackle (Hypopyrrhus pyrohypogaster), a Col- pyrohypogaster (Icteridae) Inferidas a Partir de ombian endemic and the only member of this genus, Secuencias de ADN Mitocondrial are enigmatic. Based on phylogenetic analyses of se- quences from the cytochrome b and ND2 mitochon- Resumen. Las a®nidades ®logeneÂticas de Hypopyrr- drial genes, we present a ®rst hypothesis on the sys- hus pyrohypogaster,eluÂnico miembro de este geÂnero tematic position of Hypopyrrhus. In so doing, we com- endeÂmico de Colombia, son enigmaÂticas. En este estu- plete the sampling at the genus level for a published dio establecemos la posicioÂn sistemaÂtica de Hypopyrr- phylogeny of New World blackbirds (Johnson and hus con base en anaÂlisis ®logeneÂticos de secuencias de Lanyon 1999). Hypopyrrhus belongs in a clade com- los genes mitocondriales citocromo b y ND2. AdemaÂs, posed of mostly South American species within the completamos el muestreo a nivel geneÂrico para una grackles and allies assemblage, one of the ®ve main ®logenia del grupo publicada por Johnson y Lanyon lineages in the Icteridae. Within this lineage, Hypo- (1999). Hypopyrrhus pertenece a un clado compuesto pyrrhus is consistently placed in a clade with Gym- principalmente por especies suramericanas dentro del nomystax mexicanus and Lampropsar tanagrinus, but grupo de los changos y chamones, uno de los cinco parsimony and maximum-likelihood reconstructions clados principales de los Icteridae. Dentro de este gru- do not agree on which of these species is its sister po, Hypopyrrhus se ubica consistentemente en un cla- taxon. Vocal and morphological data suggest that Hy- do junto con Gymnomystax mexicanus y Lampropsar popyrrhus and Gymnomystax are likely sister taxa. The tanagrinus, pero las reconstrucciones de parsimonia y addition of Hypopyrrhus does not cause major topo- maÂxima verosimilitud se contradicen en cuanto a cuaÂl logical changes to Johnson and Lanyon's (1999) phy- de estas especies es su taxoÂn hermano. Datos de vo- logeny of the grackles and allies; thus, the conclusions calizaciones y morfologõÂa sugieren que Hypopyrrhus y of evolutionary studies that used this phylogeny as a Gymnomystax son probablemente geÂneros hermanos. baseline remain largely unaltered. La incorporacioÂn de Hypopyrrhus a la ®logenia de Johnson y Lanyon (1989) no causo cambios topoloÂ- Key words: Andes, Colombia, Gymnomystax mex- gicos importantes, por lo que las conclusiones de los icanus, Hypopyrrhus pyrohypogaster, Icteridae, Lam- estudios evolutivos que usaron esta ®logenia como propsar tanagrinus, molecular phylogeny. base no fueron alteradas. The Red-bellied Grackle (Hypopyrrhus pyrohypogas- Manuscript received 24 September 2003; accepted ter) is one of the most poorly known New World 27 March 2004. blackbirds (Jaramillo and Burke 1999). The species is 4 E-mail: [email protected] endemic to the Colombian Andes, where it occurs lo- 5 Present address: Department of Biology, Univer- cally in the northern sector of the Central and Western sity of Puerto Rico, P.O. Box 23360, San Juan, PR Cordilleras, and in the Eastern Cordillera from De- 00931-3360. partment Caqueta south to the Ecuadoran border [664] SHORT COMMUNICATIONS 665 (Cuervo 2002). Until very recently, basic information netic hypothesis on the af®nities of Hypopyrrhus, com- on the behavior, habitat, diet, and vocalizations of pleting the sampling at the generic level for the grack- Hypopyrrhus was not available (Jaramillo and Burke les and allies phylogeny. 1999, Cuervo 2002). Although the understanding of the natural history of METHODS H. pyrohypogaster has increased substantially over re- Specimens were collected at two localities in the Mu- cent years (Cuervo 2002), its phylogenetic af®nities nicipality of Amal® (Dept. Antioquia) and deposited remain enigmatic. The species was originally de- in the collection of the Instituto de Ciencias Naturales, scribed in the genus Cassicus (De Tarragon 1847), in Universidad Nacional de Colombia (ICN). An unsexed which a wide variety of icterids including some oro- juvenile (ICN 33976) and an adult female (ICN 33977) pendolas and caciques were initially grouped. Soon af- were taken at Finca Bodega Vieja, RõÂo RiachoÂn wa- ter, Bonaparte (1850) placed it in the monotypic genus tershed, Vereda Salazar (68589N, 758039W; 1505 m), Hypopyrrhus and although a few early authors listed and an adult male (ICN 34534) at Bosque de Guaya- it under Quiscalus (Sclater 1862, Cassin 1866), bito, Finca Hugo RoldaÂn, Quebrada Caracolõ water- Hypopyrrhus was later generally accepted (e.g., Sclater shed (68519N, 758069W; 1780 m). Tissue samples taken and Salvin 1879). None of the studies of blackbird from the specimens were deposited in the collection of systematics or general revisions of the group (e.g., Instituto Alexander von Humboldt (catalog numbers Beecher 1951, Jaramillo and Burke 1999) commented IAvH-BT 2077, 2078, and 2261, respectively), togeth- on the af®nities of Hypopyrrhus, and aside from an er with blood samples drawn from each of 12 addi- unsupported statement of close relationship to the ge- tional birds that were captured and released at the same nus Macroagelaius (Ridgely and Tudor 1989), no hy- study sites and at other locations in northern Antio- potheses about its closest relative have been proposed. quia. Likewise, the position of Hypopyrrhus in linear clas- In recent years, the mitochondrial genes cytochrome si®cations (Sibley and Monroe 1990, Jaramillo and b (cyt b) and ND2 have been sequenced for the ma- Burke 1999, Dickinson 2003) is variable, an indication jority of blackbird species (Lanyon 1992, 1994, John- of the uncertainty about its af®nities. This uncertainty son and Lanyon 1999, Lanyon and Omland 1999, Om- is likely the result of the morphological dissimilarity land et al. 1999, Price and Lanyon 2002, 2004). By of Hypopyrrhus to other icterids and of the general sequencing these same two genes for Hypopyrrhus we lack of information about this taxon. There are very were able to capitalize on the preexisting database to few museum specimens in existence and most of these establish its evolutionary af®nities. We followed the are of limited scienti®c value; for example, the single laboratory protocols outlined in Johnson and Lanyon skeletal specimen for the genus available in museum (1999) to sequence a total of 1941 base pairs of collections is unsexed and incomplete (Webster 2003). mtDNA (900 of cyt b and 1041 of ND2) for two of Moreover, the present whereabouts of the type speci- the new Hypopyrrhus specimens (ICN 33977 and ICN men are unknown (Hellmayr 1937). 34534). These sequences (GenBank accession num- Hypopyrrhus is presumed to be a member of the bers AY572450±AY572453) were aligned to blackbird grackles and allies assemblage, one of ®ve main clades sequences obtained from GenBank using Sequencher in the family Icteridae (Lanyon and Omland 1999). 3.1. First, to determine whether Hypopyrrhus fell This monophyletic group comprises 42 species in 18 clearly within one of the ®ve major blackbird lineages genera (Dickinson 2003) distributed through much of identi®ed by Lanyon and Omland (1999), we did a the New World. More than half of these genera are preliminary unweighted parsimony analysis using a monotypic, which indicates a traditional lack of un- dataset containing ND2 and cyt b sequence data for 80 derstanding of relationships within the group and the blackbird taxa (see Appendix for species list and distinctiveness of many taxa. Based on mitochondrial GenBank accession numbers). The results indicated DNA sequence data, Johnson and Lanyon (1999) de- that Hypopyrrhus in fact belongs within the grackles veloped a phylogeny for the grackles and allies that and allies clade, and speci®cally within group 1 of that helped clarify relationships among genera and species assemblage as de®ned by Johnson and Lanyon (1999). and has served as a baseline for studies on the evo- This result was supported by 90% of bootstrap pseu- lution of breeding behavior, nesting habitat, plumage doreplicate data sets. An additional parsimony analysis characters, and traits presumed to be adaptations for in which transversions were weighted 2 to 1 over tran- brood parasitism (Searcy et al. 1999, Johnson and Lan- sitions produced the same result with 93% bootstrap yon 2000, Mermoz and Ornelas 2004). In their taxo- support. nomic sampling, Johnson and Lanyon (1999) included Having con®rmed that Hypopyrrhus belongs within representatives of all the grackle genera except group 1 of the grackles and allies clade, we restricted Hypopyrrhus because tissue samples of this taxon were intensive phylogenetic analyses to this assemblage, lacking at the time of their study. Thus, the phylogeny considering the 16 species for which DNA sequence of the grackles and allies is currently incomplete
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