In California, the Golden Gate National Recreation Area encompasses 80,000 acres of land smattered along the north and south entrances of the San Francisco Bay and connected by the Golden Gate Bridge. To the south of the iconic bridge are 30,000 acres of steep, rolling hills of buildings, busy streets, snuggled Victorian era houses, and, amongst it all, Golden Gate Park, the west coast’s (20% larger) equivalent of New York City’s Central Park . To the north of the bridge, away from the San Francisco metropolis, lies 2,000 acres of sagebrush-covered peninsula. Here, the Pacific Ocean grabs at the coastal cliffs, as if trying desperately to get inside the San Francisco Bay, and the ocean wind and fog are cradled by the mountains, filling up the valleys before spilling over into the sleepy harbor town of Sausalito: The Marin Headlands. 

This is a tale about the importance of community and urban connectivity; the petite and poofy main characters inhabit both sides of the Golden Gate Bridge, charismatic, with a tear-dropped plume that bobbles just over their heads, the California quail (Callipepla californica). 

The California quail is a small, nonmigratory game (i.e. a bird that is usually hunted for sport) bird that lives life low to the ground. They scan the underbrush for seeds, manzanita, poison oak berries, and occasionally insects, using dense shrubbery as a protective home for their nests and from predators [1-2]. They can fly, but prefer to run on their short and powerful legs (with a running ground speed timed at ~12 miles or 19.3km per hour) [2]. When surprised, they swiftly scuttle from open meadows and pathways towards the protection of tall grasses and shrubs, a sight that pairs well with their twittery pit-pit-pit alarm calls. 

Male and female California quails are around the same size and sexually dichromatic, meaning there is a difference in coloration between the sexes. Both males and females have that distinctive tear-drop of six feathers that droop just over their heads, but the males’ have a black coloration and females have more brown coloration. You can distinguish California quails from their closest relative (the Gambel’s quail) by the California quail’s vibrant scale-patterned chest feathers [1-2].

The Muwekma Ohlone tribe along the coast in present-day San Francisco regularly ate the plump birds as part of their regular diet. As San Francisco morphed into a cosmopolitan metropolis in the 1900s, quails maintained their presence. According to ornithologist Florence Merriam Bailey in 1902: “the brushy parts of Golden Gate Park in San Francisco abound with quail… one can watch the squads of plump hen-like little creatures as they move about with stately tread or stand talking sociably in low monosyllables” [3]. In 1931, the California quail became the state bird of (you guessed it) California and later the official bird for the city of San Francisco. Yet, despite the once plentiful squads that Bailey observed “hurry[ing] across the path like hens before a wagon” in the city’s parks and Presidio, there are no longer any California quails in San Francisco despite stable populations elsewhere (a.k.a ‘local extinction’). The last two quails seen in the city were Fajita from the Presidio in 2008 and Ishi from Golden Gate Park in 2017 [3-4]. The two solo males could occasionally be heard calling out for fellow quails with their classic chi-ca-go calls, only to be met with silence (…or more accurately, the commotion of city sounds). 

What Fajita and Ishi were missing was the gregarious, communal life that California quails thrive in. Gathering in “coveys” of up to 75 individuals with multiple, unrelated males, a few females, and many offspring, they stay together for most of the year [2,5-7]. California quails forage on the ground together, with at least one quail in the covey perched on higher ground as a lookout while the others feed [1-2]. During the breeding season, males and females may wander off from the group. Quails use a variety of mating strategies, including seasonal monogamy (1 male, 1 female), polygny (1 male, multiple females), and polyandry (1 female, multiple males) [5-7]. Females lay 12-16 eggs per clutch, sometimes laying multiple clutches with different males. Both clutches can survive because males help look after the hatchlings. So, a female can leave her hatchlings with one father, while going to lay another clutch with another father. The babies begin walking right after hatching and are raised by both parents, or sometimes communally raised, with all parents contributing to care of all offspring within the covey, regardless of genetic relation [2,6-8]. Research on California quail joint living has even shown that adults that raise young amongst other adults and offspring have a longer lifespan than adults that do not [6,8]. 

Lack of community living wasn’t the only barrier Fajita, Ishi, and past city-dwelling California quails were up against – there were literal barriers as well. To the north of hilly San Francisco lies an equally hilly space, but rather than densely urban, the Marin Headlands is densely natural, abundant with plumpy parties of birds bibbling about through the sagebrush and succulents. While other birds may pass through different green spaces across a city with little issue, California quails don’t fly much, thus requiring corridors of natural habitat that connect green spaces together. Yet increased development, fenced yards, outdoor pets, and heavy traffic made it impossible for San Francisco quails to thrive [10]. Now, several entities in the San Francisco community are exploring the idea of reintroducing the official city bird.

When doing any sort of species reintroduction, it is vital to consider the viability of a local population. Hence, researchers got to work identifying what features California quails need, not only for reintroduced birds to survive, but so future generations continue to thrive. Using data from eBird, a community science app where folks can log bird observations they see in their area, observations of California quails in various settings were compared with what qualities (e.g. size, greenery, soil type, predator presence) those green spacesparks had. Researchers found that our petite and poofy birds of interest are more likely to be found in less isolated parks with less human-made substrate (i.e. cement, asphalt, brick) and high shrub cover with open areas for foraging [9]. More quails were observed in larger parks, but once parks got larger than 3.3miles² / 5.3km² there was a high probability that local extinction would occur. This is likely because California quails are non-migratory and because they prefer to skitter across the ground (non-human made ground, that is) than fly, so dispersing across large areas is unlikely [9]. All these urban park features make sense of what we know about California quail life history and how well they thrive just north of the Golden Gate Bridge. However, one surprising feature increased the odds of a park being occupied by the quails: coyote presence! Despite quails being potential prey, these top urban predators likely suppress outdoor cat populations and other smaller urban predators (think raccoons, rats etc.) that more frequently feast on quails and their eggs [9,11]. In fact, research continues to demonstrate the detrimental impact domestic house cats have on local wildlife [4,11]. 

We are still far away from replenishing San Francisco’s green spaces with the squads of plump hen-like little creatures Florence Bailey witnessed in the 1900s and the chi-ca-go calls that Ishi and Fajita once sang. Reintroduction of California quail will not happen without the buy-in of local neighborhoods [3,4]. Humans are responsible for creating a more stable urban ecosystem; one where cats are kept indoors, corridors are a standard part of city planning, native plants and shrubs are prioritized, and new developments are built strategically to minimize environmental impact of previously natural spaces [4,10]. Until the official city bird returns to San Francisco, folks will just have to take a trip across the mouth of the bay and immerse themselves in the wonderous natural space the Marin Headlands offers for California quails and humans alike.

---

Karli Chudeau is a graduate student in the Animal Behavior Graduate Group and a part of the UC Davis Coastal and Marine Sciences Institute. She is interested in conservation management and improving animal welfare in wildlife rehabilitation settings. Her current research examines how we can use behavioral management interventions, such as environmental enrichment, to improve reintroduction success with pinnipeds. She is also an avid ocean nerd.

[Edited by Jacob Johnson and Alice Michel]

References:

[1] Calkins, J.D., Gee, J., Hagelin, J.C., & Lott, D.F. (2014). California Quail (Callipepla californica) version 2.0. In P.G. Rodewald (Ed.), The Birds of North America. Cornell Lab of Ornithology, Ithaca, New York, USA.

[2] Leopold, A. (1977). The California Quail. Berkeley, Los Angeles, California: University of California Press.

[3] Chamings, A. (2020, December 24). San Francisco’s official bird is locally extinct but may be reintroduced to the city. SF Gate, https://www.sfgate.com/local/article/SF-official-bird-is-locally-extinct-quail-15827302.php

[4] Robertson, M. (2017, October 27). There’s only 1 quail left in San Francisco, and cats are likely to blame. San Francisco Examiner, https://www.sfexaminer.com/archives/presidio-may-bring-california-quail-back-to-san-francisco/article_33fb40c9-37c1-5d0b-83ef-2bf086b02f60.html

[5] Roth, A.M., Gee, J.M. & Keiser, C.N. (2021). Autumn copulatory behavior in California quail: Observations and potential functions. Western Birds, 52, 112-113.

[6] Calkins, J.D. (2007). The family behavior of California quail: A molecular analysis. The Condor, 109, 962-967.

[7] Raitt, R.J. (1960). Breeding behavior in a population of California quail. The Condor, 62, 284-292.

[8] Lott, D.F & Mastrup, S.N.A. (1999). Facultative communal brood rearing in California quail. The Condor, 101, 678-681.

[9] Iknayan, K.J., Wheller, M.M., Safran, S.M., Young, J.S., & Spotswood, E.N. (2021). What makes urban parks good for California quail? Evaluating park suitability, species persistence, and the potential for reintroduction into a large urban national park. Journal of Applied Ethology, 59(1), 199-209, https://doi.org/10.1111/1365-2664.14045

[10] Golden Gate National Recreation Area (2021, November). Can San Francisco parks support the reintroduction of the California quail? National Park Service. https://www.nps.gov/articles/000/can-san-francisco-parks-support-the-reintroduction-of-california-quail.htm

[11] Hawkins, C.C., Grant, W.E., & Longnecker, M.T. (1999). Effect of subsidized house cats on California birds and rodents. Transactions of the Western Section of the Wildlife Society, 35, 29-33.

All pictures taken by the author unless otherwise specified.