Gene D. Sattler

MORPHOMETRIC VARIATION AS AN INDICATOR OF GENETIC INTERACTIONS BETWEEN BLACK-CAPPED AND CAROLINA CHICKADEES AT A CONTACT ZONE IN THE APPALACHIAN MOUNTAINS

Abstract We studied hybridization and introgression between Black-capped (Poecile atricapillus) and Carolina (P. carolinensis) chickadees along two transects in the Appalachians using four genetic markers and multivariate analysis of morphology. Genetic data revealed that at least 58% of the birds in the center of each transect were of mixed ancestry and that recombinant genotypes predominated among hybrids, demonstrating that hybridization is frequent and that many hybrids are fertile. Genetic clines generally were steep and coincident in position, but introgression was evident well beyond the range interface. Introgression was higher at the one autosomal locus surveyed than in mitochondrial DNA or in two sex-linked markers, suggesting that the hybrid zone is a conduit for gene flow between the two forms at some loci. On a broad scale, morphometric variation was concordant with genetic variation. Clines in morphological variation based on principal components (PC) scores were steep and coincident with genetic clines. Also, a strong correlation within a population between PC scores and an individuals genetic makeup suggested that a large amount of morphological variation was genetically determined. However, morphological analysis indicated that hybrids were uncommon on one transect, whereas genetic data clearly showed that they were common on both. In addition, patterns of morphological variation were equivocal regarding introgression across the hybrid zone. Thus, genetic data provided a complementary and more detailed assessment of hybridization, largely due to the discrete nature of genetic variation. Genetic markers are useful in understanding hybridization and introgression, but diagnostic markers may underestimate average gene flow if selection against hybrids maintains steep clines at diagnostic loci. To gain a clearer picture of the genome-wide effects of hybridization, a much larger number of loci must be assayed, including non-diagnostic ones.

REPRODUCTIVE SUCCESS ACROSS THE BLACK-CAPPED CHICKADEE (POECILE ATRICAPILLUS) AND CAROLINA CHICKADEE (P. CAROLINENSIS) HYBRID ZONE IN OHIO

Abstract Black-capped Chickadees (Poecile atricapillus) and Carolina Chickadees (P. carolinensis) hybridize in an east-west band from New Jersey to Kansas. Within the past century, the Ohio portion of this hybrid zone 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 evidence of assortative mating in the center of the hybrid zone, we found a relative paucity 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. Éxito Reproductivo a través de la Zona de Hibridación de Poecile atricapillus y P. carolinensis en Ohio

AN ASSESSMENT OF SONG ADMIXTURE AS AN INDICATOR OF HYBRIDIZATION IN BLACK-CAPPED CHICKADEES (POECILE ATRICAPILLUS) AND CAROLINA CHICKADEES (P. CAROLINENSIS)

Abstract Vocal admixture often occurs where differentiated populations or species of birds meet. This may entail song sympatry, bilingually singing birds, and songs with intermediate or atypical characteristics. Different levels of vocal admixture at the range interface between Black-capped Chickadees (Poecile atricapillus) and Carolina Chickadees (P. carolinensis) have been interpreted as indicating that hybridization is frequent at some locations but not others. However, song ontogeny in these birds has a strong nongenetic component, so that inferences regarding hybridization based on vocal admixture require confirmation. We used diagnostic genetic markers and quantitative analyses of song to characterize population samples along two transects of the chickadee contact zone in the Appalachian Mountains. More than 50% of individuals at the range interface were of hybrid ancestry, yet only 20% were observed to be bilingual or to sing atypical songs. Principal component analysis revealed minimal song intermediacy. This result contrasts with an earlier analysis of the hybrid zone in Missouri that found considerable song intermediacy. Re-analysis of the Missouri data confirmed this difference. Correlation between an individual’s genetic composition and its song type was weak in Appalachian hybrid populations, and genetic introgression in both forms extended far beyond the limits of vocal admixture. Therefore, song is not a reliable indicator of levels of hybridization or genetic introgression at this contact zone. Varying ecological factors may play a role in producing variable levels of song admixture in different regions of the range interface. Una Evaluación de la Mixtura de Cantos como Indicador de Hibridación en Poecile atricapillus y P. carolinensis

An assessment of song admixture and its reliability as an indicator of genetic ancestry in Black-capped and Carolina Chickadees

Abstract Vocal admixture often occurs where differentiated populations or species of birds meet. This may entail song sympatry, bilingually singing birds, and songs with intermediate or atypical characteristics. Different levels of vocal admixture at the range interface between Black-capped Chickadees (Poecile atricapillus) and Carolina Chickadees (P. carolinensis) have been interpreted as indicating that hybridization is frequent at some locations but not others. However, song ontogeny in these birds has a strong nongenetic component, so that inferences regarding hybridization based on vocal admixture require confirmation. We used diagnostic genetic markers and quantitative analyses of song to characterize population samples along two transects of the chickadee contact zone in the Appalachian Mountains. More than 50% of individuals at the range interface were of hybrid ancestry, yet only 20% were observed to be bilingual or to sing atypical songs. Principal component analysis revealed minimal song intermediacy. This result contrasts with an earlier analysis of the hybrid zone in Missouri that found considerable song intermediacy. Re-analysis of the Missouri data confirmed this difference. Correlation between an individual’s genetic composition and its song type was weak in Appalachian hybrid populations, and genetic introgression in both forms extended far beyond the limits of vocal admixture. Therefore, song is not a reliable indicator of levels of hybridization or genetic introgression at this contact zone. Varying ecological factors may play a role in producing variable levels of song admixture in different regions of the range interface. Una Evaluación de la Mixtura de Cantos como Indicador de Hibridación en Poecile atricapillus y P. carolinensis

The effect of aerodynamic braking on the inertial power requirement of flapping flight: case study of a gull

There has been an unresolved question of whether there is any significant degree of aerodynamic braking during wing deceleration in the flapping flight of birds, with direct analogies existing with flapping micro air vehicles. Some authors have assumed a complete conversion of kinetic energy into (useful) aerodynamic work during wing deceleration. Other authors have assumed no aerodynamic braking. The different assumptions have led to predictions of inertial power requirements in birds differing by a factor of 2. Our work is the first to model the aerodynamic braking forces on the wing during wing deceleration. A model has been developed that integrates the aerodynamic forces along the length of the wing and also throughout the wing beat cycle. A ring-billed gull was used in a case study and an adult specimen was used to gather morphometric data including a steady state measurement of the lift coefficient. The model estimates that there is a 50% conversion of kinetic energy into useful aerodynamic work during wing deceleration for minimum power speed. This aerodynamic braking reduces the inertial power requirement from 11.3% to 8.5% of the total power. The analysis shows that energy conversion is sensitive to wing inertia, amplitude of flapping, lift coefficient and wing length. The aerodynamic braking in flapping micro air vehicles can be maximised by maximising flap angle, maximising wing length (for a given inertia), minimising inertia and maximising lift coefficient.

The Energy Benefits of the Pantograph Wing Mechanism in Flapping Flight: Case Study of a Gull

Bird wings generally contain a 4-bar pantograph mechanism in the forearm that enables the wrist joint to be actuated from the elbow joint thus reducing the number of wing muscles and hence reducing the wing inertia and inertial drag. In this paper we develop a theoretical model of inertial power for flapping flight to estimate the advantage of the 4-bar pantograph mechanism by comparing the inertial power required for the case where wrist muscles are present in the forearm with the case where wrist muscles are not present in the forearm. It is difficult to predict how wrist muscles would look when there is no pantograph mechanism. Therefore a lower bound and upper bound case are defined. The lower bound case involves redistributing the elbow muscles with no increase in wing mass. The upper bound case involves replicating the biceps-triceps muscles near the wrist joint. At minimum power speed the model estimates that the 4-bar pantograph mechanism reduces the inertial power for the gull from between 6.1%–12.3% and reduces the overall power by 0.6%–1.2%. When account is taken of the tight margins involved in the design of a flying vehicle, the energy savings produced by the pantograph mechanism are significant. A ring-billed gull was chosen for the case study and an adult specimen was obtained to gather morphometric data. Lessons for the design of flapping micro air vehicles are discussed.