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And Carolina Chickadee (P Liberty University DigitalCommons@Liberty University Faculty Publications and Presentations Department of Biology and Chemistry 2005 Reproductive Success Across the Black-capped Chickadee (Poecile atricapillus) and Carolina Chickadee (P. carolinensis) Hybrid Zone in Ohio C. L. Bronson Thomas C. Grubb, Jr. Gene D. Sattler Liberty University, [email protected] Michael J. Braun Follow this and additional works at: http://digitalcommons.liberty.edu/bio_chem_fac_pubs Recommended Citation Bronson, C. L.; Grubb, Jr., Thomas C.; Sattler, Gene D.; and Braun, Michael J., "Reproductive Success Across the Black-capped Chickadee (Poecile atricapillus) and Carolina Chickadee (P. carolinensis) Hybrid Zone in Ohio" (2005). Faculty Publications and Presentations. Paper 35. http://digitalcommons.liberty.edu/bio_chem_fac_pubs/35 This Article is brought to you for free and open access by the Department of Biology and Chemistry at DigitalCommons@Liberty University. It has been accepted for inclusion in Faculty Publications and Presentations by an authorized administrator of DigitalCommons@Liberty University. For more information, please contact [email protected]. The Aiik 122(3):759-772, 2005 © The Americiin Ornithologists' Union, 2005. ^L Printed in USA. REPRODUCTIVE SUCCESS ACROSS THE BLACK-CAPPED CHICKADEE {POECILE ATRICAPILLUS) AND CAROLINA CHICKADEE (P. CAROLINENSJS) HYBRID ZONE IN OHIO C. L. BRONSON,' ' THOMAS C. GRUBB, JR./ GENE D. SATTLER,' AND MICHAEL J. 'Dqmrtiticnt of Evolution. Ecohi^y, and Organlsmal Biology, The Ohio Staff Uiiivcrsifii, 318 West Tllli AveiUK Columims, Ohio 43210. USA: 'Di-piut)Jiciit of Biolo;^y mid Cliemistri/, Libert]/ Uniz'crsity, 1971 University Boiilcvnrii, i\/mhbuy^, Vir^iniiJ 24502, USA; mid iciit ofVcrldinUe Zoolo;^y, Nntioiuil Miisciofi ofNntiirnl History, Siiiillisoniiin Institution, 4210 Silver tliil Roud. Siiiliand, Mmyhmd 20746, USA CT. —Black-capped Chickadees (/'(ifc;7fi7fr;L"(7/)/7/i/s) and Carolina Chickadees (P. cawliueusis) hybridize in an east-west band from New Jersey to Kansas. Within the past century, the Ohio portion of this hybrid /one and the Carolina Chickadee range to the south have been moving northward, whereas the Black-capped Chickadee range has retracted. In Ohio, we characterized the genetic composition of the hybrid zone using five diagnostic molecular loci. Although there was no evi- dence of assortative mating in the center of the hybrid zone, we found a relative pau- city of genetically intermediate breeding females as compared with breeding males. That suggests viability selection against female hybrids, in line with Haldane's rule. On the basis of reproductive variables (number of nestlings, reproductive success), we found a decrease in productivity of breeding pairs in the hybrid zone that is significantly and positively related to their probability of producing homozygous offspring at each autosomal or sex-linked locus. We also found that the decrease in productivity was significantly and positively related to the genetic composition of the male of the pair {i.e. pure male chickadees more productive). These data strongly suggest that hybrids are at a selective disadvantage. Because the zone of reduced reproductive success was considerably narrower than the zone of introgression, our results demonstrate that genetic introgression is occurring in the face of substantial selection against hybrids. Received 16 April 2004, accepted W jmniary 2005. Key words: Black-capped Chickadee, Carolina Chickadee, genetic indices, hybrid zone, Poecile iitricapillus, Poecile carolinensis, reproductive success. Exito Reproductivo a traves de la Zona de Hibridacion de Poecile ntricapillus y P. carotiiicnsis en Ohio RESUMEN. —Las especies Poecile atricapillus y P. caroliucnsis hibridan en una franja orientada de este a oeste desde New jersey hasta Kansas. Durante el ultimo siglo, la seccion de Ohio de esta zona de hibridacion y el rango de P. carolinensis al sur de esta se han desplazado hacia el norte, mientras que ei rango de P. atricapillus se ha contraido. En este estudio, caracterizamos la composicion genetica de la zona de hibridacion en Ohio usando cinco loci moleculares diagnosticos. Aunque no cxistio evidencia de apareamiento asociativo en el centro de la zona de hibridacion. 'Present address: Department of Internal Medicine, Division of Immunology, The Ohio State University, 473 West Twelfth Avenue, Columbus, Ohio 43210, USA. E-mail; bronson.3("osu.odti 759 760 BRON-ION Lr AL. [Auk, Vol. 122 encontramos una relativa carencia de hembras reproducHvas geneticamente intermedias en comparacion con los machos reproductivos. Esto sugiere la existencia de seleccion por viabilidad en contra de las hetnbras hibridas, lo que concuerda con la regla de Haldane. Con base en variables reproductivas (nt'imero de pichones, exito reproductivo), encontramos una disminucion en la productividad de las parejas en la zona de hibridacion que esta significativa y po.sitivamente rclacionada con SLi probabilidad de producir crias homocigoticas en cada locus autosomico o ligado a! sexo. Tambien encontramos que la disminucion en la productividad estuvo significativa y positivamertte relacionada con la composicion genetica del macho de la pareja (i.e. los machos puros fueron mas prodiictivos). Estos datos sugieren fuertemente que los hibridos se encuentran en desventaja seloctiva. Debido a que la zona de exito reproductivo reducido fue considerablemente mas estrecha que la zona de introgresion, nuestros resultados demuestran que a pesar de que existe seleccion en contra de los hibridos, esta sucediendo introgresion genetica. ZONE ij-iNAMics is a fertile area for -100 km farther north, approximately along U.S. research on natural selection and speciation, Highway 30 (Grubb et al. 1994, Peterjohn 2001). because of the exchange of genes between In other words, the Black-capped Chickadee distinct groups (Harrison 1990, 1993; Hewitt distrihtition has been receding northward. 1988). Within birds (see review in Grant and The chickadee hybrid zone is quite narrow, Grant 1992), the hybridization of many North with genetic cline widths on the order of 20 American species has been studied (for review to 30 km (Sattler 1996, Sattler and Braun 2000). of Great Plains hybrid zones, see Rising 1983). Given the likely age of contact and the dispersal For example, Black-Capped Chickadees (Poecile capabilities ot chickadees (Weise and Meyer atricapillus) and Carolina Chickadees {P. caroli- 1979), the narrow cline widths suggest that ncnsis) are known to hybridize in many areas some sort of selection may oppose introgression along their common border (e.g. Kansas: across their hybrid zone {Barton and Gale 1993). Rising 1968; Missouri: Braun and Robbins 1986, In Illinois, Brewer (1963) found that hatching Sawaya 1990; Illinois: Brewer 1963; Ohio: Grubb success was lower in the chickadee hybrid zone et al. 1994; Virginia: Johnston 1971, Sattler 1996, than for either parental species, but he had Sattler and Braun 2000; West Virginia: Sattler complete data on only four hybrid zone nests. 1996, Sattler and Braun 2000; Pennsylvania: He attributed the reduction to infertility and Ward and Ward 1974, Cornell 2001). Because retarded development of eggs. these species may not be sister taxa (Gill et al. We wished to study the relationship between 1989, 1993; but see Sattler and Braun 2000 for hybridization and reproductive success in greater discussion), only limited hybridization might detail. The objectives here were to employ genetic be expected. markers to map one segment of the hybrid zone In North America, except for a peninsular in north-central Ohio and to examine the rela- distribution in the Appalachian MoLintains, tionship between genetic composition of the the Black-capped Chickadee distribution abuts parents and reproductive success. the north edge of the Carolina Chickadee di.s- tribution (Mostrom ct al. 2002). In the southern Appalachians, Black-capped Chickadees are often found at high elevations, and Carolina Field nicthods. —The area of the hybrid zone Chickadees at lower elevations. In the early studied within Ashland County, Ohio (4O''5O'N, 1880s, Carolina Chickadees were described as 82"15'W) was bounded by County Road 700 on permanent residents only within the southern the north. State Route 95 on the south. State portion of Ohio (Wheaton 1882). By the late Route 89 on the west, and County Road 175 on 1930s, the hybrid /one was probably located the east (Fig. 1). The study area was 23 km from acrt)ss the middle of the state, approximately in north to south and 6 km from west to east. The the location of the east-west U.S. Interstate 70 landscape was about equally divided among (Trautman 1940). Currently, the zone is located pasture, row crops, and woodlands. To limit the July 2005] clive Success in tli/hriii Zone 761 overlap of points in Figures 2-4, the 21 sampling In November of 1993 and 1994, we placed locations were condensed to 10 pooled s^imples remote-controlled Feeder traps (Pierce and based on similar latitudes. Frt>m north to south, Grubb 1979) filled with sunflower seeds within the groupings were a-b, c, d, e, g-f, h, i-m, n-p, privately owned woodlands af all study sites q-r, and s-u (Fig. 1). within the zone. From December through CR700 5- 10- km 15- 25- FIG. 1. Tbe study transect (light gray area) in Ashland County, Ohio, indicating sampling points (letters) and major roadways. 762 BKONSON hi' AL. [Auk, Vol. 122 February, we trapped or mist-netted chickadees and Pst l/mtDNA) into a genetic index (Cl) visiting each feeder. In late February, we placed for each individual, calculated as tbe number artificial nesting snags (Grubb and Bronson of Carolina Chickadee alleles divided by tbe 1995) in the woodlands and monitored them total number of alleles examined (Bronson et al. through the chickadee breeding season (to the 2003). Tbere are two alleles for eacb autosomal end of June). marker and one for tbe mtDNA haplotype. Tbe At the time of capture, we banded each bird Z-linked marker has two alleles in males and with a federal aluminum band and a colored one in females (females are tbe heterogametic leg streamer (Sullivan 1984) tor individual sex in birds).
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