Relationship between Flight Feathers: Primary Feathers, Tail Feathers, and Ecological Considerations

Kalvin Foo Department of Biology The College of New Jersey Ewing, NJ 08628 Foo2@tcnj.edu

ABSTRACT Flight is a function primarily powered by a birds wings, but in order to control for speed, brake, and steer, the bird is reliant on its tail. This study utilized a sample of American Robins (Turdis Migratorius) to compare the lengths of tail feathers to primary feathers in conjunction to a longitudinal study of tail feather lengths in order to ascertain growth factors. A positive correlation was found between primary and tail feathers (N=25, r= .75, p < .05) that suggested tail feathers being a limiting factor for primary feather growth (m=.86). No ecological stressors were able to be determined from the longitudinal study that would directionally select for tail sizes.

INTRODUCTION From their evolutionary origin as flightless dinosaurs, flight-capable birds have evolved to have optimal physiologies to enhance their survivability and efficiency in the air. While the focus of many studies regarding airborne travel is the role of the wing and its feathers, the tail and its feathers play a significant role in bird flight. Most tail morphologies are primarily influenced by aerodynamic considerations of the species to optimally serve flight, with the exception of species where sexual selection chooses for aerodynamically unfavorable tails (Thomas, 1997). For flight-capable birds, their tails must then work in conjunction with the lift and drag forces already generated by their wings in order to help maneuverability in terms of steering, braking, and speed control. The inner and outer wings of birds are known for creating lift and thrust, respectively, which can be modified by changing the angle of attack. Studies have shown that, despite the small amount of lift that tails do contribute during flight, they create more drag than if the bird had a shorter tail, reducing the birds lift to drag ratio (Thomas & Balmford, 1995). Selective pressures thus dictate the birds tail shape in order to maintain function without creating too much drag. With respect to this, tail usage is optimized at slower speeds, as the stability and lift it produces reduce the power necessary at for slow flight (Thomas, 1997). As such,

morphological data from a significant sample size comparing the birds outer wing, as determined from the primary feathers, to its tail length, as determined from the tail feathers, should show that a positive correlation between primary and tail feather lengths. Correlation between the two feathers should give insight into the impact of their relationship on bird flight. Because great influence in prior research focuses on the negative effects of the tail, this study was performed hypothesizing that tail feathers in amongst a species will be selected to be shorter. The study made use of American Robins (Turdis migratorius), a year-round residential species to the area where the study was performed, Southern New Jersey. Longitudinal data over time could give implications towards presence of selective pressures over time in the area. This is based off of a theory presented by Price, Chi, Pavelka, and Hack (1991) suggesting that morphological variances in feather size in a population stem from varied conditions during childhood. Presence of mounting or dissipating ecological pressure would show in tail lengths, as need for more maneuverability or greater speed would correspond with increased or decreased tail lengths.

METHOD Twenty-five preserved American Robins (Turdis migratorius) collected from general South Jersey locations were temporarily acquired from Princeton University with assistance of Dr. L. Butler. Specimens ranged in age from 1878 to 2010, and were recorded by previously issued identification numbers. Measurements were performed with calipers capable of measuring accurately to a tenth of a millimeter. Primary feathers were measured on the left wing, as viewed from a ventral view, from the most anterior point of bend (wrist) to the furthest posterior point of

the primary projections. Tail feathers were measured from the point at the end of the specimens vent down the tip of the left tail feather, on the same side of the measured wing. Experimenter bias was removed via blind data collection means, as prior collected data on the same specimen was obscured during second and third measurements. Measurement repeatability was validated by finding a mean coefficient of variation (V), as determined from the means and standard deviations from three measurements of each specimen Statistical analysis was conducted via Statistica, ver. 7, using the basic correlation analysis utility.

RESULTS Primary feather length was found to average 119.7 mm (N= 25, V= 2.6%) and the average tail feather was 81.7 mm (N=25, V= 6.2%). Primary feathers ranged from 105.9 mm to 129.5 mm (Fig. 2) Notably, there are two outliers in tail feather length from the general range of lengths from 74.8 mm to 87.1 mm, each with high coefficients of variation (V > 9.2%). Statistical analysis determined no correlation between feather lengths and time (p> 0.05; Fig. 1). Positive correlation was determined between primary feather and tail feather lengths (p < 0.05; Fig 2). The two variables correlated at r= 0.75, and the calculated correlation line was found to have a slope value of 0.86.

DISCUSSION Significant ecological implications could not be drawn from a longitudinal correlation analysis, as year distribution from samples was highly modal and was not conducive to finding a statistically significant correlation. However, this implicates that there were no specific growing or declining selective pressures for tail feather lengths. Deforestation studies in southern New

Jersey have shown that the majority species affected are neo-tropical migratory birds- decreasing their survivability odds (Rich, Dobkin, & Niles 1994). From this, a resident species such as Turdis migratorius may have sufficient maneuverability ability to survive unimpeded by changes in habitat structure. This supported by the wide variety of habitats American Robins have been found in, as reported by the Sibley Guide to Birds. Conversely, the wide distribution in different tail feather lengths per year (Fig. 1) childhood conditions of the sample specimens imply highly varied childhood conditions (Price et. al, 1991). From this, it can be drawn that although no directional selection is occurring over time, individuals are still presented with differing conditions that are affecting tail feather formation. Positive correlation between tail feathers and primary feather length was found as expected, for at a basic level larger individuals should have longer feathers. As there was a significant correlation found (Fig. 2), a point of note was the slope value of the best fit line (m=.86). Because the slope is less than 1, it implies that relative to increase of tail length feathers, primary wing feathers increase less. On the other hand, it could also be seen as a need for a higher rate of tail growth to compensate for increased primary feather length. This data suggests that longer primary feathers are able to compensate for the drag produced by a longer tail, and that a longer tail may be necessary to steer and brake against the greater forces from the wings. In a species, the morphological adherence to this correlation occurs within boundaries, as too large or too small a feature will be inefficient and unfavorable. Data showed that tail feathers had a smaller range of sizes; potentially supporting that primary feather length may be contingent on limits of tail size, as wings too large may produce forces too great to be efficiently managed by a limited tail size.

This study focused on Turdis migratorius in order to focus on variations within a singular species as per environmental fluxes instead of differences between species. However, a study conducted across a family of species may give further insight into genetic and evolutionary bases behind the flight relationship between birds tails and outer wings.

LITERATURE CITED

Price, T., Chi, E., Pavelka, M., Hack, M. 1991. Population and developmental variation in the feather tip. Evolution 45: 518-533. Rich, A., Dobkin, D., Niles, L. 1994. Defining forest fragmentation by corridor width: The influence of narrow forest-dividing corridors on forest-nesting birds in Southern New Jersey. Conservation Biology 8:1109-1121. Thomas, A.L. 1997. On the tails of birds. Bioscience 47:215-225. Thomas, A.L., Balmford, A. 1995. How natural selection shapes birds tails. The American Naturalist 6:848-868.

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Tail Feather Length (mm)

86 84 82 80 78 76 74 72 70 68 1860

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Figure 1. Correlation of Tail Feather Lengths and Year. Tail feather lengths of Turdis migratorius were plotted with respect to year. Correlation failed the 95% confidence interval (p= 0.92)

132 130 128 126

Primary Feather Length (mm)

124 122 120 118 116 114 112 110 108 106 104 68 70 72 74 76 78 80 82 84 86 88 90 92 94

Tail Feather Length (mm)

Figure 2. Correlation of Tail Feather and Primary Feather Lengths. Tail feather and primary feather lengths of Turdis migratorius were correlated to find a positive correlation (p =0.00), with r= .75 and line slope m= 0.86.