There are three genetic forms of male Ruff shorebirds, each with distinct plumage and breeding behaviors: independent males defend territories and have colorful plumage; satellite males have ornamented plumage but do not hold territories; female-mimicking faeder males also do not hold territories but are tolerated on independent male territories. These three forms stably coexist throughout the Ruff's breeding range.

1. 3. The Ruff is a European shorebird where males come in three different genetic forms, each
with a distinct plumage pattern, breeding strategy, and suite of behavioral characteristics.
Independent males have highly ornamented and colorful plumage patterns; they defend small
territories to which they try to attract females with elaborate plumage displays (see a video here).
Satellite males are also ornamented but are less colorful than independent males. They do not
defend territories; instead, they lurk on the periphery of the territories of Independent males and
attempt to copulate with females attracted by the Independents. Faeder males are smaller and
mimic females in body size, plumage pattern, and behavior. Like the Satellite males, they do not
defend Ruff (Philomachus pugnax) display territories; however, because of their female-like
appearance and behavior, they are tolerated on the territories of Independent males and, like the
Satellites, attempt to copulate with females attracted to the Independents. All three genetic forms
coexist in stable polymorphism throughout their breeding range. A. Consider two gene loci in the
Ruff, one contributing to variation in male plumage pattern and one contributing to variation in
male territorial behavior. Would you expect the alleles at these two loci to be in linkage
equilibrium or linkage disequilibrium? Briefly explain (4 points). B. Based on what you learned
in class about similar complex genetic polymorphisms in Heliconius butterflies and White-
throated Sparrows, what predictions would you make about the genetic architecture and
proximity of these two gene loci within the Ruff genome? Briefly explain (4 points).