New analysis out of the Hubbard Brook Experimental Forest reveals that migratory songbirds use their songs to equally allocate breeding territories amongst themselves.
From the Spring 2026 problem of Residing Chook journal. Subscribe now.
It’s early June within the White Mountains of New Hampshire, a transparent, cool, calm morning—about an hour after daybreak. The rising solar and a greening forest create an ever-shifting dapple of sunshine and shade throughout the forest ground.
Carrying her cellphone in a single hand, and a pair of binoculars round her neck, Abigail Archey—a rising junior majoring in organic sciences at Cornell College—strides in straight traces via the trailless wilderness. The uneven terrain is crammed with rotting logs, ankle-catching swales, and clusters of shin-whacking hobblebush. As she traverses the forest, she rattles off chicken species she hears: “Crimson-eyed Vireo. Wooden Thrush. Black-throated Inexperienced. Black-throated Blue.”
Just like the tons of of scientists earlier than her who’ve walked this very same floor on the Hubbard Brook Experimental Forest for the previous 57 years, Archey is making historical past, one chicken at a time. She data every chicken on an ArcGIS mapping app on her cellphone. Some songs are barely audible, some clear and shut. However there’s barely any motion to be seen. Almost each chicken is recognized by sound alone.
“In the event that they weren’t singing, we wouldn’t know most of them have been right here,” she says.
“Right here” is Hubbard Brook. The experimental forest was began by the U.S. Forest Service in 1955 in partnership with a number of Dartmouth School professors, establishing the 7,800-acre northern New England woodland as a spot to review the long-term results of forest administration. Within the many years that adopted, Hubbard Brook turned a multidisciplinary hub of scientists finding out practically each side of ecology and biology inside the forest’s northern hardwood ecosystem, from snails to soil vitamins to snowpack. And, famously, birds.
The oblong Hubbard Brook “chicken plot,” as scientists right here name it, occupies simply 10 hectares (round 25 acres, or about 0.3% of the forest’s whole space), nevertheless it has performed an outsized position in serving to researchers perceive the breeding season habits of migratory songbirds. That’s as a result of there’s a file of practically each breeding chicken that has hung out on this plot since 1969. The information comes from practically six many years of researchers surveying this slice of forest on the top of songbird breeding season, recording each chicken they see or hear and noting the places and actions to be able to get correct counts of what number of birds are current.
There’s nothing prefer it in all of ornithology, in response to Mike Webster, a professor of neurobiology and habits at Cornell College.
“I can’t consider some other examples the place you discover this a lot data over this a lot time for this many species,” says Webster, who additionally oversees the Cornell Lab of Ornithology’s Hubbard Brook Field Ornithology Program, which brings in undergraduates like Abigail Archey every summer season to achieve area expertise and help with analysis.
“Mapping particular person territories was actually only a option to rely the birds,” says Richard Holmes, who was simply two years into his profession as a Dartmouth professor in 1969 when he began the Hubbard Brook chicken plot. “The maps weren’t the primary endeavor. They have been a software to assist us get correct numbers of breeding birds.”
Now because of a brand new digitization mission, these previous maps are yielding shocking new insights about how communities of migratory songbirds form their territories on their breeding grounds. In a research revealed within the journal Ecology Letters in 2024, scientists analyzed these maps to see how the territory sizes for various songbirds within the chicken plot have modified from season to season. It seems the birds keep bigger territories when there are fewer people within the plot, however can shrink their territories to accommodate extra people within the plot when populations at Hubbard Brook go up—offering proof that these warblers and different forest birds behave in response to a concept generally known as very best free distribution.

Scott Sillett, who leads the Smithsonian Migratory Chook Heart, has studied the habits of songbirds like warblers and thrushes for the reason that mid-Nineties when he obtained his PhD below Holmes at Dartmouth. He says this analysis units a brand new bar for understanding the underlying dynamics of how songbirds set up their breeding territories.
“It’s a type of research you type of pinch your self, like, ‘wow! we’re capable of present a very sturdy check of type of a foundational concept in ecology,’” he says.
Sillett additionally says that it’s notable the research relied on a vital side of those songbirds—particularly, their songs—to measure and observe how they set up their territories.
“Tune is a option to be aggressive with out bodily attacking your neighbor,” he says. “It’s less expensive. I imply, when you’re an organism with hole bones, how a lot precise bashing into one other animal do you need to endure?”

A 12 months within the life of a newly fledged Black-throated Blue Warbler at Hubbard Brook performs out like an motion film. It begins late within the spring, when the younger chicken hero casts out from the nest by itself. Come autumn, the chicken travels midway throughout an unknown world, to the tropics and again, going through obstacles reminiscent of buildings, extreme climate, and restricted stopover habitat for resting and refueling. It perseveres, striving to return to the place the journey started, its birthplace, to finish its quest—profitable replica, the possibility to convey a brand new era into the world.
In line with the long-term knowledge from Hubbard Brook, there may be a dozen or extra Black-throated Blue Warblers seeking to breed on the chicken plot every year, in addition to a pair dozen Crimson-eyed Vireos, and anyplace from a number of to a horde of birds of at the very least a dozen different species. How can all of those breeding birds—generally effectively over 100 people in all—discover a breeding associate and set themselves up with the assets wanted to attain their very own Hollywood endings?
Mike Webster—who can also be director of the Cornell Lab’s Macaulay Library, the world’s largest scientific archive of pure historical past audio, video, and images—has studied chicken communication in songbirds world wide, from the warblers at Hubbard Brook to Crimson-backed Fairywrens in Australia. Webster says vocal communication, reminiscent of songs throughout breeding season, play an enormous position in giving every chicken a preventing probability in a crowded enjoying area.
“Migratory birds want to ascertain a territory rapidly, earlier than different returning people lay declare to the most effective breeding habitat,” he says. “Songs permit these birds to right away promote their declare on unfilled habitat … they’ll simply broadcast that ‘this space is mine.’”
In line with Webster, birdsong is the proper medium for staking a declare to breeding territory in forested habitat, as a result of track travels effectively via the timber—in contrast to visible indicators reminiscent of brilliant plumage, which might be obscured in dense forest foliage.
“With out songs, it’s laborious to think about that birds would have the ability to be environment friendly sufficient to breed efficiently within the small window they need to be up right here,” says Webster, noting that Black-throated Blue Warblers are solely at Hubbard Brook from Could to August. “Songs are just like the glue that maintain chicken societies collectively.”
Richard Holmes, now a professor emeritus on the cusp of his ninetieth birthday, has been listening to the “zu-zu-zu-ZHWEE” of the black-throated blues at Hubbard Brook since he first flagged off the chicken plot. As of 2025, Holmes was nonetheless making his option to the sphere station at Hubbard Brook to advise, observe, and take part in knowledge assortment. He says that though the chicken songs sound like peaceable melodies to people, they’re truly pugnacious warnings about staking out turf.
“A lot of the aggression we see happens shortly after arrival, simply earlier than and through egg-laying,” he says. For half-ounce, 5-inch-long Black-throated Blue Warblers, that aggression takes the type of frequent chases, shut encounters, and sometimes, bodily assaults. Holmes says that inside a few weeks, the breeding territory boundaries are inclined to settle out and singing turns into the birds’ important approach of sustaining their declared areas.
From the start of his research at Hubbard Brook, Holmes established data-gathering methods, subsequently utilized by a number of generations of scientists and scientists-in-training, to find out what number of breeding birds are occupying the plot. Key to the success is what Holmes and his collaborators dubbed “wander mapping.” That’s what Abigail Archey was doing on that early June morning, and it’s the important thing to creating positive researchers aren’t lacking any birds that may be current. It’s a way that entails strolling the plot every morning for a complete of 16 hours per week, for 5 weeks straight, on the top of chicken breeding season.

“‘Wander mapping’ was simply what we began calling it. Actually it needs to be referred to as ‘territory mapping,’” says Holmes. In contrast to straight-line, timed transects (that are additionally performed via the plot), wander mapping provides the researcher company to maneuver freely to allow them to “cowl all components of the plot, and to comply with particular person birds if wanted,” Holmes says. “They’ll find yourself discovering nests, noting chases and fights between males, and documenting the presence of females, all of which add to the accuracy of our counts.”
Maybe most significantly, the wander-mapping researcher should decide when two or extra males of the identical species are countersinging. Holmes says that countersinging is usually detected when one male is singing and a number of of the close by birds are reacting in sort.
“So many instances when one chicken sings, one other chicken responds,” says Holmes. “It’s like these birds are having … a territorial rap battle. When you’ve gotten a number of data of this countersinging habits, you’ll be able to see, okay, they’re shifting nearer collectively, they’re farther aside, and you may draw a boundary about midway in between these birds.”
For greater than 50 years, every chicken detection and occasion of countersinging was plotted on paper with a grid representing the chicken plot. Every birdsong will get a dot (with a time) and an episode of countersinging is indicated by a line with arrows connecting two of the dots. On the finish of every season, the researchers would use translucent paper to switch all of the each day maps for one species onto a single map for the season. Because the researchers analyzed the info for every species, the dense thickets of dots and arrows reworked into an interlocking sequence of round blobs, every representing a territory.
Starting in 2023, the chicken territory mapping mission at Hubbard Brook went absolutely digital, with a smartphone app that enables every researcher to enter knowledge instantly on to a shared map. Miranda Zammarelli helped with the digital transition as a PhD scholar at Dartmouth. Right this moment she oversees the summer season fieldwork and knowledge assortment at Hubbard Brook. She says that regardless of the technical upgrades, the method for figuring out the boundaries of chicken territories stays a lot the identical because it was when Holmes began it.
“I have a look at the info for the season, and I make the boundaries myself attempting to match as shut as doable to the paper approach,” Zammarelli says. “That is the best way that they’d been doing it for 50 years,” she says.



When Zammarelli arrived at Dartmouth in the summertime of 2021, she instantly began brainstorming tasks for her dissertation. Her advisor tipped her off in regards to the 50-plus years of paper map information saved away in cupboards at Hubbard Brook—practically 1,000 of them.
“I spent two weeks type of zoning out on the scanner, interested by the questions we may reply with these knowledge,” says Zammarelli. “For me there was type of the ‘a-ha’ second of seeing all of those maps go by and really with the ability to see a number of the adjustments in circle sizes over time simply by trying them over.”
Zammarelli observed the territories could be larger in some years and smaller in others. That led her to provide you with the concept to check a long-standing pair of theories about how birds of the identical species occupy habitats as an space will get extra crowded.
One of many theories—generally known as very best free distribution, or IFD—holds that animals will unfold throughout habitats to equalize their means to outlive and reproduce. Meaning particular person territory sizes will lower in proportion to one another when the inhabitants will increase; in consequence the animals divide up the out there habitat roughly equally.
A competing concept—very best despotic distribution, or IDD—holds that extra dominant people monopolize high-quality areas, forcing others into poorer habitats, which might result in the territories in the most effective habitats staying bigger whilst the encircling inhabitants will increase. Whereas these theories had been examined in particular person species, with various outcomes, nobody had examined them with a number of species over the timescale of many years.
Zammarelli started by specializing in the territories of seven of the traditionally most considerable breeding songbird species on the chicken plot: Black-throated Inexperienced Warbler, Black-throated Blue Warbler, American Redstart, Crimson-eyed Vireo, Ovenbird, Hermit Thrush, and Least Flycatcher.
Along with the newly scanned territory maps, she additionally used long-term knowledge in regards to the total abundance of every of those species—which years have been increase years, which have been bust—in addition to further data gleaned from the maps about which areas have been the preferred locations to arrange a territory.


“We made these warmth maps that confirmed us which areas have been essentially the most most well-liked for every species to nest in, all-time. We use that as a sign of which areas within the plot are the upper high quality spots,” says Zammarelli. For instance, Black-throated Blue Warblers are more likely to cluster in areas of the plot with stands of hobblebush, the place practically all of their nests are discovered.
Zammarelli says that understanding habitat high quality is vital to figuring out if the birds are free to pick out their territory, or in the event that they’re topic to potential despots of their midst.
“If these birds have been being despots, we might see that birds which are nesting in essentially the most most well-liked areas would keep the identical measurement areas from 12 months to 12 months,” she says. “The concept is, when you’re in the highest quality habitat, you’re not going to change your territory measurement simply because extra people are coming in. You’re the dominant chicken, you’re going to carry on to that house and pressure everybody else into the low-quality habitat.”
Nonetheless, Zammarelli’s evaluation confirmed that songbird societies at Hubbard Brook forest have a tendency towards the egalitarian. Throughout all of the species she checked out, territory measurement decreased as native inhabitants elevated—so extra birds may match into the house. She says the truth that birds averted being habitat hogs on the extra most well-liked websites makes a robust case that they function in an IFD method, a end result that shocked Scott Sillett, the Smithsonian Migratory Chook Heart chief.
“What shocked me was … we all know that these warblers are territorial, they’re aggressive, they dive-bomb one another…. That’s why I feel loads of us anticipated it to be very best despotic distribution, like, ‘effectively, after all, they’re little despots!’” says Sillett.
Zammarelli factors to an instance from the Ovenbird maps to point out the way it performs out.
In 2001, the researchers tallied round seven Ovenbirds on the 25-acre plot, one of many lowest counts of the entire research. That contrasts with 2009 when round 21 Ovenbirds, thrice as many, crammed into the identical space.In line with the calculations from the paper, the typical measurement of every particular person territory in 2001 was round 5 acres, whereas the typical territory measurement in 2009, when there have been many extra birds, was about 2.5 acres.
“You may as well see a rise in countersinging within the 12 months when there are extra people,” she says. “Theoretically, you probably have extra neighbors, you’re in all probability going to need to sing extra to defend your house.”

Zammarelli stresses that this is only one instance, and that countersinging charges weren’t a part of their evaluation within the analysis revealed in Ecology Letters.
Regardless of the dip and surge within the early 2000s, Ovenbirds have maintained a secure inhabitants at Hubbard Brook since round 2010, says Zammarelli, because of rising areas with mature timber and sparse understory—the popular breeding habitat for the species. In line with Richard Holmes, Hubbard Brook is now not the younger forest it was when he arrived within the late Nineteen Sixties.
“The forest was logged closely via in regards to the 1910s,” he says. “After I first got here right here it was nonetheless thought of early successional, fewer previous timber.”
Two of essentially the most considerable birds from the primary twenty years of Hubbard Brook chicken surveys—American Redstart and Least Flycatcher—desire youthful forests, and so they have stopped nesting on the plot fully because the forest has matured. Populations of different species—reminiscent of Black-throated Inexperienced Warbler and Crimson-eyed Vireo, which desire extra mature forests—have proven a slight however regular constructive development, although there can nonetheless be big variations from 12 months to 12 months. For instance, Crimson-eyed Vireos on the plot have numbered as few as 13 in 1980, and as many as 32 in 2016.
Holmes is making ready to submit his most up-to-date evaluation of the Hubbard Brook chicken abundance tendencies in a scientific journal, including to the greater than 140 journal articles revealed utilizing Hubbard Brook chicken knowledge on which he has been an creator. He says that whereas the general inhabitants of songbirds on the plot decreased by round 60% between 1970 and 2000, the entire songbird inhabitants on the plot has trended barely upwards over the previous twenty years.
Zammarelli, who accepted a place as a postdoctoral researcher on the Cornell Lab beginning within the fall of 2026, says that whereas there isn’t a established relationship between both of the distribution theories and inhabitants tendencies, the truth that the general inhabitants of birds is at the very least holding regular, and probably rising, is encouraging. She says that the birds’ tendency towards very best free distribution might be a constructive factor for the best way these birds can climate adjustments to habitat and meals availability.
“With very best free distribution, birds ought to capable of reproduce fairly equally on high- and low-quality habitats,” says Zammarelli. “So, if one thing occurs to a part of the habitat, there ought to nonetheless be people who can achieve reproducing. With despotic distribution, [birds] in the highest quality habitat may be the one ones which are reproducing efficiently.”
Zammarelli additionally factors out that in the case of conserving forests like Hubbard Brook for birds, this research factors to the worth of including less-than-ideal habitat to panorama conservation. The outcomes of her analysis present that these songbird species have the power to distribute themselves in a approach that gives alternatives for equal health throughout patchworks of high-quality and low-quality habitat.



“We shouldn’t simply focus all our efforts on the high-quality areas. We’d like to consider the lower-quality areas, as a result of these areas nonetheless equal some reproductive output for birds,” she says.
“These birds have fairly a little bit of flexibility,” she says, “which has implications for the way they’ll climate local weather change or habitat loss.”
Mike Webster, the Cornell Lab’s Hubbard Brook Subject Ornithology program director, says that these sorts of insights and conclusions about chicken inhabitants dynamics are solely doable from long-term research just like the practically 60 years of knowledge gathered at Hubbard Brook.
“[This study] is known as a testomony to the facility of long-term research,” says Webster. “To know how birds reply to local weather change and different environmental disturbances over time, it is advisable to return to the identical spot, year-after-year, and accumulate these knowledge. There may be simply no different approach of doing it.”
