It鈥檚 One of the Biggest Events in a Bird鈥檚 Life. Scientists Can鈥檛 Agree How to Talk About It.

Every species replaces its feathers by molting, but a passionate debate among ornithologists about what to call the plumage swaps spans oceans and generations.
Portrait of a mottled brown Rock Ptarmigan looking straight on at the camera.
Rock Ptarmigan. Photo: Debra Herst/爆料公社 Photography Awards

Feathers define birds as we know them. Whether fanned in courtship display, slicing the air, or gliding through water, birds鈥 plumage makes them unique among living animals. But as fascinating and glorious as feathers may be, they are merely dead tissue, and any damage to them is permanent. Too many impaired feathers spells disaster for a bird. Luckily, birds can replace their mangled plumes鈥攁nd they do so at least once a year in a process called molting, often right under the noses of birders. Some species take the opportunity to sport seasonal looks, like the nearly radioactive red of a male Scarlet Tanager in spring. But most importantly, molt is a survival tactic, necessary to keep feathers in peak condition. Simply put: If a bird doesn鈥檛 molt, it will die.

When, where, and how often a bird replaces its plumes varies widely between and even within species, making molt a formidable research subject, though one that is key to understanding a species鈥 life history and conservation needs. Compared to other major events in birds鈥 lives, namely breeding and migration, molt is understudied. 鈥淚t鈥檚 amazing,鈥 says Erik Johnson, director of conservation science at and author of a book on the molt of neotropical birds, 鈥淭here are so many ornithologists that just don鈥檛 think about molt.鈥 Yet it鈥檚 a vital part of all avian lives: 鈥淣ot all birds breed every year. Not all birds migrate. But every single bird molts every year. It鈥檚 that foundational to what a bird is.鈥 

But ornithologists and molt-curious birders are likely to encounter a conundrum when diving into the subject: A schism divides the world of molt studies, with a major border line running down the Atlantic Ocean. Compare a scientific paper on molt from Europe to one written in the Americas, and you may be at a loss to find their common ground (and not only because Europeans spell it as 鈥渕oult鈥). The two camps tend to use completely different words to talk about molt. 

A schism divides the world of molt studies, with a major border line running down the Atlantic Ocean.

While the system favored in Europe is tied to other events in birds鈥 lives鈥攂reeding, in particular鈥攖he terminology used in the Americas was designed by ornithologists Philip Humphrey and Kenneth Parkes in the 1950s to refer solely to molting itself. So, for example, when a tanager is turning scarlet, a European ornithologist would likely call it a 鈥減re-breeding molt,鈥 which results in a 鈥渂reeding plumage.鈥 An American using the Humprhey-Parkes system would deem the same transition a 鈥減realternate molt,鈥 yielding the tanagers鈥 鈥渁lternate鈥 plumage. (Further complicating the picture, many bird banders around the world, including in North America, use a separate, system to describe age based on molt and the calendar year in which a bird hatched.)

At first glance, the main difference between the competing terminologies seems to be that the European strategy, sometimes called the 鈥渓ife-cycle approach,鈥 is intuitive and easily mapped onto a bird鈥檚 life, while the Humphrey-Parkes (H-P) system is bewilderingly abstract: A freshly fledged bird is in its first basic plumage, sometimes quickly followed by a preformative molt, and perhaps later by a first prealternate molt鈥攁ll within its first year of life. Even H-P proponents admit the terminology can be confounding, and years of conflated ideas and disagreements about how to apply the terms led to a to the system in 2003.

So why bother with a confusing, unintuitive way to talk about an already complicated process? Indeed, a 2022 paper to get on the same page and adopt the 鈥渃learer and more understandable鈥 life-cycle terminology, arguing that the simpler language could even enable recreational birders to understand molt and contribute to its study.

The life-cycle approach to molt, though, has a major problem. The system works well for the birds it was developed to describe: the small songbirds frequently banded and studied across Europe. By and large these birds replace all their feathers in the fall, after breeding (the 鈥減ost-breeding鈥 molt), and many molt again in the spring, some into flamboyant colors useful for attracting a mate (the 鈥減re-breeding,鈥 or, in older works, 鈥減re-nuptial鈥 molt). But things quickly get messy moving beyond northern songbirds. Raptors and large waterbirds don鈥檛 fit the pattern very neatly, and the system really starts to break down in the tropics, where birds鈥 lives are shaped by very different pressures and resources. Many tropical birds lack predictable breeding seasons and can reproduce throughout the calendar year, even interrupting their molt to do so. Some species undergo several molts in their first year, which the life-cycle approach struggles to classify. It鈥檚 not just a few outliers, either. Because of the in the tropics, the life-cycle system doesn鈥檛 work well for the majority of the world鈥檚 bird species.

鈥淭he terminology gets complex because the biology is complex, not just because we want to complicate our lives,鈥 says Miguel Moreno Palacios, an ornithologist at Universidad de Ibagu茅 in Colombia. As a young ornithologist, training with scientists from the United States and Europe, Moreno Palacios learned quickly that much of what he was taught from northern systems would need tweaking鈥攐r replacing. He鈥檚 not compelled by the argument that it鈥檚 easier to grasp a 鈥渟impler鈥 way of talking about molt, even if it doesn鈥檛 hold up in large parts of the world. 鈥淭hat鈥檚 not a good reason,鈥 he says. 鈥淭hat鈥檚 not how science works.鈥

__________

Because of the overwhelming concentration of bird diversity in the tropics, the life-cycle system doesn鈥檛 work well for the majority of the world鈥檚 bird species.

__________

Humphrey and Parkes attempted to head off these problems 65 years ago, with a fresh they hoped would 鈥渢hrow new light鈥 on the study of molt, which they argued was being held back by the muddling of concepts. As one example they offered the Rock Ptarmigan, found in the arctic and alpine regions of several European countries (and in North America across the arctic tundra). In Scotland, Humphrey and Parkes reported, the bird breeds in its brown 鈥渟ummer plumage.鈥 Meanwhile in Greenland the same species breeds with snowy white 鈥渨inter鈥 feathers. (And in Scandinavia ptarmigans breed in 鈥減articoloured, partimoulted dress.鈥) 鈥淥bviously,鈥 the pair wrote, 鈥渋n this species no one plumage may correctly be called a 鈥榥uptial鈥 plumage in the functional sense.鈥

Humphrey and Parkes had likely seen for themselves the shortcomings of older molt terminology, first in their work with museum specimens from the tropics, and later in the field. As curators of natural history collections, they were taxonomists used to thinking about lineage, and besides accounting for the world鈥檚 many 鈥渁berrant鈥 birds, their proposal had a deeper mission: to map molt onto the evolution of birds as a class of animals. Considered this way, the H-P system is simpler than a 鈥渓ife-cycle鈥 approach that needs myriad exceptions and qualifications to work for every bird around the world. Humphrey and Parkes instead looked to the 鈥渇undamental pattern鈥 they saw across all species of birds and went from there. The different ways birds molt today, they asserted, can be seen as clues to the branches that have split and sprouted from the avian tree of life over millions of years.

More than half a century later, H-P boosters are still inspired by that vision. 鈥淚 don鈥檛 think they even fully understood how great their system was,鈥 says Peter Pyle, a biologist at the Institute for Bird Populations and the author of a pair of exceedingly detailed that opened the door to molt for many North American ornithologists, bird banders, and even enthusiastic birders (Pyle also helped developed universal , based on the H-P system).

Earlier this year Pyle and co-authors published a that foregrounds and builds on Humphrey and Parkes鈥檚 evolutionary perspective. An illustration shows how (the authors hypothesize) different molt strategies evolved along different lineages of birds. Pyle hopes that homing in on molt as an evolved, and evolving, trait can help make the system feel more intuitive. The evolutionary lens also calls attention to thought-provoking details鈥攍ike the increasingly clear evidence that molt is a carry-over from ancestral reptiles shedding their skin. Similarly, the appearance of extra molts in many distantly related groups of birds suggests the strategy evolved several different times in avian history. These molts, called prealternate in the H-P system, are the ones that some birds now take as an opportunity to don more brilliant plumes, useful for attracting a mate. Since so many of these more-frequent molters are long-distance migrants or live in harsh environments, lifestyles that are hard on feathers, it seems likely that this strategy evolved first and foremost as a needed rejuvenation. Flashy courting outfits are just a bonus鈥攄early appreciated by birders.

If your head is spinning from the dueling molt terminology, there鈥檚 no need for distress, says Johnson. Instead of worrying too much about which system to use, he recommends just tuning in to molt itself. When and where do your local birds molt? Can you see it in action? Learning about molt, he says, will make you a better birder and add to your appreciation of the birds around you and the challenges they face. But if you want to go deeper on the subject, definitely stick with Humphrey-Parkes, Johnson says, because you can use it anywhere in the world: 鈥淚t鈥檚 a system that is inherently global." 

Given the life-cycle approach鈥檚 lapses in the tropics and for many groups of larger birds, it鈥檚 unclear why the entire field of ornithology hasn't embraced the H-P system鈥攖hough Pyle does have a somewhat less-than-scientific theory: 鈥淓uropeans hate it when Americans come up with something better than they have.鈥

This year鈥檚 paper was Pyle鈥檚 latest attempt to persuade his colleagues across the Atlantic to 鈥渟ee the light,鈥 he says. 鈥淲e鈥檒l see if it works.鈥