The Red Quail Mystery

Jul 18, 2022

Red Quail, Revisited: How Herbert Stoddard’s Specimens are Helping Solve a Centuries-Old Bobwhite Mystery

If, like me, you have ever found yourself scrolling through the Bobwhite Quail Breeders Facebook group (for purely academic reasons, I assure you), then perhaps you are already familiar with so-called “Tennessee Red” bobwhites. True to their name, both males and females of this gorgeous color morph sport all over auburn red plumage and a characteristic white patch in the center of their breast, and are among the most popular variety of bobwhites in the commercial market (along with other fantastic breeds such as “snowflakes,” “Mexican speckleds,” and “Wisconsin jumbos”). Unlike many of these other varieties, however, red bobwhites are known to occur occasionally in the wild (one was taken by a hunter in the Red Hills region last year). Now commonly used to train bird dogs, red bobwhites have a history that far predates our current era of online quail shopping and were, in fact, the object of much fascination and scientific study by Herbert Stoddard himself at Tall Timbers nearly 90 years ago.

Gould’s painting of Ortyx castanea

Gould’s painting of Ortyx castanea from his 1850 publication A Monograph of Odontophorinae, or Partridges of America. Although Gould originally thought this was a new species from South America, it is now considered to be the first record of the rare red morph of Northern Bobwhites. Image in the public domain

Bobwhite Comparison

One of the original red bobwhites sent to Stoddard from Ames (left); a bobwhite from an aviary showing intermediate “reddish” plumage; a typical bobwhite from Louisiana (right). Specimens were photographed in the Louisiana State University Museum of Natural Science. Photo by Jessie Salter

History of Red Bobwhites
Stoddard compiled an excellent history of red bobwhites, which he first described in his seminal work The Bobwhite Quail: Its Habits, Preservation, and Increase in 1931, and later expanded upon in a 1949 publication in the ornithological journal The Auk. Stoddard traced the first description of this rare color morph to British ornithologist John Gould, who purchased a bobwhite with distinctive red plumage from a market in Manchester, England and described it as a new species in 1842. Gould named this species Ortyx castanea (“castanea” being the Latin root for chestnut), and although he originally believed it to be a previously undescribed species from South America, he later amended that it could instead be a rare mutation of a typical bobwhite from the United States.

The next record of a red bobwhite does not appear until 1921, when a single female with this unusual coloring was shot in King County, Virginia, but in 1927 reports of multiple coveys containing red birds began surfacing from the Ames Plantation in Grand Junction, Tennessee. The manager of Ames wrote to Stoddard in 1927, describing these birds and later published an article about them in The American Field (thus inspiring the name “Tennessee Reds”), which introduced red bobwhites to a much wider audience of hunters and ornithologists alike. Stoddard visited Ames in 1928 to see these red quail for himself, and described them as looking “almost brilliant” in direct sunlight. Capitalizing on this newfound public interest, Ames began a captive breeding and release program for their red bobwhites.

The timing of the rediscovery of red bobwhites coincided with a wave of genetic studies on domestic bird plumages in the early twentieth century. Just a few years earlier, geneticists at the University of Wisconsin published a study describing the inheritance pattern of a similar red color morph in domestic pigeons, and in 1930 several red quail from Ames were sent to the University of Wisconsin to repeat the experiments with bobwhites. The quail proved somewhat less cooperative than the pigeons, however, and the results of the study were stymied by the poor survival of the offspring.

Many of the parents and offspring from Stoddard’s 1930s breeding experiments are preserved as museum specimens, along with his hand-written notes, in the Herbert Stoddard Bird Collection at Tall Timbers. Photo by Jessie Salter

Stoddard’s Experiments on Red Quail
In 1934, Ames arranged to have the study repeated at Sherwood Plantation near Thomasville, Georgia under the guidance of Stoddard, who had established himself as the authority on bobwhites. Stoddard and his colleagues designed an elegant series of breeding experiments using red birds from Ames and bobwhites with typical plumage from the Red Hills region. Between 1934-1936, they performed 14 crosses and recorded the plumage of the resulting offspring.

Through these experiments, Stoddard and his colleagues discovered that when red bobwhites were bred with normal bobwhites their offspring had an intermediate, “reddish” plumage, a pattern known as incomplete dominance. They also demonstrated that unlike typical bobwhites, of which males and females can be distinguished by plumage, males and females with red plumage were seemingly identical.

These findings were significant because red plumage in pigeons was found to be a sex-linked trait, indicating there were multiple genetic pathways through which red plumage could be produced in birds. Stoddard and his colleagues eventually published the results of their study in The Auk in 1949, but despite the rise in popularity of red bobwhites among commercial breeders in the following decades, it appears their findings were largely forgotten by the broader scientific community.

I first learned about Stoddard’s interest in red bobwhites when I visited Tall Timbers for the first time in 2018. I had come to spend a few weeks working with and learning from the Game Bird Lab during their January trapping season. As a museum-based ornithologist who is often the only person in the room studying bobwhites, it was invigorating to be immersed in a community of fellow bobwhite enthusiasts who were kind enough to share their tremendous knowledge with me. One of the game bird lab members asked if I’d ever read Stoddard’s study on red bobwhites (referring to the 1949 article in The Auk); I had not, but I read it that day and was immediately intrigued.

Over the past two decades, advances in molecular techniques and computational analyses have facilitated a new wave of fascinating studies into the genetic basis of bird coloration, led by pioneering studies in species like pigeons, chickens, and Coturnix quails that have historically been the targets of selective breeding for plumage traits. This is no coincidence; the knowledge of the genetic basis of these traits provided by classic studies of human-associated species is invaluable in designing effective modern studies that in turn provide insight into the evolution of plumage color across the bird Tree of Life.

In the past few years, several studies have built upon centuries of breeder knowledge to describe in detail the genes and molecular pathways involved in producing numerous plumage mutations in pigeons, including the genes responsible for sex-linked red plumage. I wondered why no one had followed up on Stoddard’s study on red bobwhites. There were still unanswered questions: Was Stoddard’s hypothesis of incomplete dominance correct? What genes were involved in producing red plumage? And what about the red bobwhites being bred in captivity today — do all red bobwhites share the same genetic mutation, or has red plumage evolved multiple times independently?

Herbert Stoddard Bird Collection
During that visit to Tall Timbers, I had carved out an afternoon to explore the Herbert Stoddard Bird Collection, which comprises nearly 4,000 bird specimens, many of which were collected and prepared by Stoddard himself. The collection is a testament to Stoddard’s exceptional commitment to documenting and preserving the natural history of the Red Hills region. While digging through drawers of bobwhite specimens I made a fantastic discovery: here were the very red bobwhites Stoddard and his colleagues described in their 1949 paper! They had prepared many of the parents and offspring from the crosses performed during 1934-1936 as specimens, and preserved them in the bird collection at Tall Timbers, along with Stoddard’s hand-written notes describing the experiments. I immediately knew what a unique and valuable resource this was.

A lot has changed in the nearly 90 years since Stoddard performed his experiments with red bobwhites. We can use technology like spectrophotometers — devices that measure the wavelengths of light transmitted by an object — to objectively measure the color of feathers and identify the pigments they contain. By taking tiny pieces of skin from the toes of historical specimens — a technique that wasn’t available even a decade ago — we can sequence the entire genomes of birds and identify the genes involved in producing different plumage colors. By using these modern methods on Stoddard’s historical specimens, we can build upon the knowledge Stoddard contributed in his original study, and continue his pursuit to understand the evolution of these remarkable red quail.

New Genome Sequencing Techniques Confirm Stoddard’s Hypothesis
Together with a team of scientists from Tall Timbers and Louisiana State University, we were able to use these new techniques to sequence the genomes of 34 historical specimens of parents and offspring from Stoddard’s original 1930s breeding experiments, including two of the red bobwhites sent to Stoddard from Ames. These historical specimens include bobwhites with red, intermediate “reddish”, and typical plumage. We’re not quite ready to publish our findings, but our results confirm Stoddard’s hypothesis that red plumage is an incomplete dominant trait in bobwhites and suggest that the genes responsible for producing red plumage in bobwhites are not the same genes that produce red plumage in pigeons, but they are known from color morphs in chickens, Coturnix quail, and other species of passerine birds. Importantly, our results also suggest that different populations of red bobwhites produce red plumage through different genetic mutations. We’re excited about these results and what they tell us about how color mutations evolve in bobwhites and other species, and we look forward to sharing our full findings in the near future!

A selection of bobwhite specimens with plumage mutations

A selection of bobwhite specimens with plumage mutations (including a Tennessee red, far right) in the Herbert Stoddard Bird Collection at Tall Timbers. Photo by Jessie Salter

Stoddard could not have known in 1934 that his specimens would one day have their entire genomes sequenced — the double helix structure of DNA wasn’t even discovered until 1953! But, he knew the importance of museum collections for preserving a record of biodiversity, and he had the foresight to ensure that future generations of scientists could build upon his tremendous contributions to the fields of natural history, ecology, and ornithology.

Also preserved in the Stoddard Bird Collection are bobwhites with other unusual plumage mutations — like albinism, melanism, and one incredibly peculiar specimen with protruding discs of feathers on each cheek — that are still waiting to be studied. Some of these birds Stoddard collected himself, and others were sent to him by hunters and quail managers across the country who knew he would be interested in studying and preserving them. Nearly a century after he began his own observations, Stoddard’s legacy as a naturalist and collector continues to shape and inspire our understanding of bobwhite biology.

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