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Karyotypes of Six Species of North American Blackbirds (Icteridae: Passeriformes)

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Karyotypes of Six Species of North American Blackbirds (Icteridae: Passeriformes) KARYOTYPES OF SIX SPECIES OF NORTH AMERICAN BLACKBIRDS (ICTERIDAE: PASSERIFORMES) HOLLYH. HOBART,1 SCOTTJ. Gu•q•q,2 A•qVJoH•q W. BICKI4AM Departmentof Wildlife and FisheriesSciences, Texas A&M University, CollegeStation, Texas77843 USA ABSTRACT.--Karyotypesof six speciesof blackbirdswere prepared using in vivo tech- niques.Five speciesare karyotypicallysimilar and may be more closelyrelated than tradi- tional morphologicalcharacters suggest. The EasternMeadowlark (Sturnella magna), how- ever, is very differentin karyotypeand may be more distantlyrelated to the other species than has been previouslythought. Karyotypicevidence is consistentwith Beecher'shy- pothesisof a closerelationship between emberizine finches and blackbirds.Received 25 June 1981, accepted18 December1981. THE family Icteridae (Passeriformes)is a di- mosomalvariation that can revealevolutionary verse group of predominantly tropical New and phylogeneticrelationships. Among the World songbirdsarranged by various investi- karyotypicallybest known vertebrategroups gators into 89-101 species in 25-36 genera are the mammalian orders Chiroptera, Pri- (Storer 1960, 1971; Blake 1968; Short 1968, 1969; mates, and Rodentia and the reptilian order Clements 1978). Beecher (1951) believed that Testudines.These groups show diverse pat- the radiation of the icteridswas an "explosive" terns of variation in the chromosomecomple- event, the family quickly filling all available ment (Capanna and Civitelli 1970, Gorman niches. Later, Beecher (1953) described Icteri- 1973,Matthey 1973,Wilson et al. 1975,Bush et dae as a terminal taxon, anatomicallyand be- al. 1977, Bickham and Baker 1979, Bickham haviorally more complexthan the group from 1981). The mammalianorders exhibit highly which it arose. variable karyotypes, while lower vertebrate Early phylogeneticstudies of the icterids taxa usually are karyotypicallyconservative. were confinedto plumagecoloration and gross Comparedto the karyologyof mammalsand morphological characters of the bill, skull, reptiles, avian karyology is relatively unex- wing, and foot. In 1951, Beecherestablished plored,with lessthan 3% of all specieskaryo- the mostrecent phylogeny of the group, based typed (Takagi and Sasaki 1974, Shields in on jaw and skull musculatureand bill shape. press).Many avian taxa show little or no vari- He hypothesizedthat the icteridsevolved from ation in standardkaryotypes, yet someavian the emberizinefinches, through the primitive genera (e.g. lunco) are composedof species genus Molothrus,in three phyletic lines: the that share identical chromosomepolymor- Agelaiine, Quiscaline, and Cassicine.Based on phisms(Shields 1973, 1976). Shields (in press) myologicaland foragingstudies, Raikow (1978) has reviewed the amount of variation in chro- believed that Emberizinaeand Icteridae may mosomesof birds, and studiescomparing the be sistergroups. karyotypesof closelyrelated bird speciesare Recently, comparative karyological studies rare. The availability of six speciesof icterids have been used to discern patterns of chro- in our area provided an opportunity for com- parative studies. There are two basic strategiesfollowed in karyotypicinvestigations. The firstis to survey • Presentaddress: Department of Ecologyand Evo- relatedforms throughout a taxonin an attempt lutionary Biology,The University of Arizona, Tuc- to determine patternsof karyotype variation son, Arizona 85721 USA. amongsubgroups. The secondis to surveya 2 Presentaddress: Department of Biology, Texas large number of individualsfrom the rangeof A&M University, CollegeStation, Texas77843 USA. a single speciesor speciesgroup to discern 514 The Auk 99: 514-518.July 1982 JULY1982] BlackbirdKaryotypes 515 Fig.1. Brown-headedCowbirdkaryotype. Acomplete chromosome setfrom a male; inset are the sex chromosomesfrom a female. croscope.Chromosomes werearranged according to whatpatterns of intraspecificvariation and size;terminology for centromereposition follows polymorphismarepresent in naturalpopula- Patton (1967). tions. At least10 metaphasocells of eachspecies except Thepurpose of thispaper is to presentthe I. graduacaudawereexamined todetermine thedip- resultsof our karyotypicstudy of these'six 1oidnumber. Both sexes were examined for each speciesrepresenting fivegenera of Icteridae: the speciesexcept I. graduacauda(male only) and Q. Brown-headedCowbird (Molothrusater), the quiscula(female only). Sturnella magna (Brazos Coun- Great-tailedGrackle (Quigcalus mexicanus), the ty,Texas) and I. graduacauda(Webb County, Texas) CommonGrackle (Q. quiscula),the Red- werecollected during the breeding season; all other individualswere trappedfrom wintering popula- wingedBlackbird (Agelaius phoeniceus), the tionsin BrazosCounty. Fifteen individuals of Black-headedOriole (Icterus graduacauda), and aterwere examined, as were at leasttwo each of the the EasternMeadowlark (Sturnella magna). otherspecies except I. graduacauda. Thisrepresents the only cytogenetic informa- Voucherspecimens deposited in theTexas Coop- tionreported for mostof thesespecies, al- erativeWildlife Collection(TCWC) at TexasA&M thoughDNA amounts were reported for Age- Universityare: NI. aterTCWC 11029, 11030, 11031, laiusphoeniceus and Sturnellamagna by 11032,10926; A. phoeniceusTCWC 11033, 11034; Q. Bachmannet al. (1972),and the karyotypeof mexicanusTCWC 11035,11036; Q. quisculaTCWC the formerspecies was reported by Makino 11028;and I. graduacaudaTCWC 10910; 10 specimens and Baldwin (1954). of M. ater,2 specimensof S. magna,1 of Q. mexi- canus,2 of Q. quiscula,and 1 of A. phoeniceusare METHODS uncatalogued. Somaticmetaphase chromosomes were prepared RESULTS from 27 icteridsusing in vivo bone marrow proce- dures.Individuals were injectedintraperitoneally MOLOTnRUS AT•R (2n = 78--80:Fig. 1), Qms- with a 0.05% solutionof colchicineat 0.3-0.4 ml per CALUSt•EXlCAt•US (2n = 76-78: Fig. 2A), Q. 100g body weight and were sacrificed after a period QCnSCULA(2n = 76: Fig. 2B),AGELAIUS PnO•NI- of 20-30min. The femurswere removed and flushed C•US(2n = 80: Fig. 2C), ICTERUSGRADIL4CAUDA witha hypotonicsolution of 0.075M KCIto obtain (2n = 76-78:Fig. 2D) a suspensionof bone marrow cells, which was in- cubatedin thehypotonic solution at 37øCfor 30 min. Thechromosomal complements of these five The cellswere then fixed (3 partsmethanol to 1 part speciesare similar. The six largest autosomal aceticacid) for 10 min and resuspended two or three pairs(macrochromosomes) aredistinguishable timesin freshfixative to ensurefixation. Slides were bysize from the remaining 31-33 pairs (micro- preparedbydropping three or four drops of cell sus- chromosomes).The largest pair of macrochro- pensionon cleanedglass slides and igniting the mosomesis submetacentric,the nextfour pairs preparation.The slides were then stained for 5 min in a warm 10% Giemsasolution. Preparations were aresubtelocentric, and the sixthlargest pair is observedand photographed on a LeitzDialux mi- acrocentric.Microchromosomes appear to be 516 HOBART,GUNN, AND BICKHAM [Auk,Vol. 09 zw Fig.2. Partialkaryotypes offive icterids. Only the 10 largest autosomes andsex chromosomes areshown forA. Great-tailed Grackle, female; B.Common Grackle, female; C.Red-winged Blackbird, female; D.Black- headedOriole, male; and E. EasternMeadowlark, female. acrocentricexcept for two pairs of biarmed centric,and the fifth autosomeis acrocentric; chromosomes.Microchromosomes arefigured thus,there are fourpairs of autosomalbiarmed onlyfor M. ater(Fig. 1). Thelack of resolution macrochromosomesas compared to five in the due to smallsize and apparentsimilarity other species.The Z chromosomeis a medium- among species limits the usefulnessof micro- sizedmetacentric, and the W is probablyac- chromosomesfor comparativestudies. We rocentric.There are 32 pairs of microchromo- haveattempted only to enumeratethem. The somes. Z chromosome,which is about the fourth larg- est chromosomein size, is subtelocentricfor DISCUSSION all of these species.The W chromosomeis probably an acrocentricmicrochromosome in Beecher(1951) hypothesized that the icterids all exceptM. ater,in whichit probablyis a evolved from the emberizine finches via the small metacentric chromosome. Makino and genusMolothrus. He describedthree phyletic Baldwin(1954) reported karyotypes for A. linesof Icteridaedescending from Molothrus: phoeniceusand Xanthocephalus xanthocephalus the Agelaiine,containing Agelaius, Sturnella, (theYellow-headed Blackbird) with macrochro- Icterus,and severalgenera not reportedhere; mosomal complementsidentical to those re- the Quiscaline,containing the subgenera portedin this study. Quiscalus(Q. quiscula)and Cassidix(Q. mexi- canus)of thegenus Quiscalus and several gen- STURNELLAMAGNA (2n = 78:Fig. 2E) era not reportedhere; and the Cassicine,none of which is reportedhere. Therefore, Molothrus Thekaryotype of S. magnadiffers from those mightbe expectedto possessa primitive,or describedfor theabove five species in several ancestral,icterid karyotype. respects.The largestautosomal element is sub- In general,the six species show little karyo- telocentric, the fourth autosome is subrecta- typicvariation. The diploidnumbers range JvL¾1982] BlackbirdKaryotypes 517 from 76 to 80 due to interspecificdifferences in for a primitive biarmed W chromosome(the the number of microchromosomes.Five species only character in which M. ater differs from appear to have identical macrochromosomal most other icterids). Certain biochemical stud- complements (Figs. 1 and 2A-D). Sturnella ies, however, indicate that Beecher was correct magnadiffers
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