What if we had an early warning system in loons that could alert us to population decline, like the proverbial canary in a coalmine?

Male loons might serve as such an early warning system. That is, careful monitoring of the health of male loons might provide a good indication of the health of the loon population as a whole. How is this possible? Because the more we study the breeding ecology of loons, the more stark differences we find between the sexes. And — more to the point — male loons have some chinks in their armor that females do not.

Most fundamentally, males are 25% larger than females. Greater size places greater energetic demands on males. Males are living “closer to the edge” than females and might often fail to acquire enough food during the season to maintain good body condition. Thus, a decrease in the quality or quantity of food — which could set in motion a population decline — should strike males first and hardest. Indeed, as the graph below shows, the average mass of male loons has declined in northern Wisconsin over the past 30 years in a way that suggests they are having more time finding food now than they used to. (Note that females have not declined in mass during the same period.) The obvious conclusion: something in Wisconsin lakes has changed in the past three decades that has impaired males’ ability to feed themselves.

Average masses of male and female loons in northern Wisconsin, 1991 to 2021. Male mass has declined significantly during this period, while female mass is unchanged.

Long before I discovered that male masses were in decline, I had begun to worry about male loons. You see, male loons live shorter lives than females. This means that there are simply fewer adult males around. In fact, the majority of non-territorial adults (“floaters”) in the loon population are females. Since males are in short supply, the loss of an adult male breeder on a lake or territory sometimes leads to that territory becoming vacant. In fact, in 23 of 24 well-documented instances where an adult breeder’s death was associated with a territory vacancy, the dead breeder was a male. Vacant territories are, of course, a harbinger of overall population decline.

Sadly, recreational fishing does not help the situation. Possibly because males’ greater size makes them a bit more desperate to feed themselves, male loons are twice as likely as females to be hooked by anglers or become entangled in fishing line. This pattern is well-documented in New England loons, but the same scenario plays out in the Upper Midwest. Specifically, of 47 known fishing entanglements among our study animals, 33 involved males, and only 14 involved females. Angling mortality, then, exacerbates what is already a female-skewed sex ratio owing to early male senescence.

It is difficult to predict the future, but I think you can see why I am concerned. Male loons appear to be in trouble. We cannot say for certain whether mass loss by male loons will cease or continue. Furthermore, we have no evidence to date that the 4% net loss in mass by males since 1991 has negatively affected their survival. So it is too early to panic about these patterns. But it is also hard not to feel like a miner glancing anxiously at his lethargic canary.

I do not think of myself as a cheerful bearer of bad news. Yet I repeatedly bring it. Each time I meet a new lake resident and secure permission to cross their land and observe their loons, I brace myself for the inevitable question: “Is it true that loons mate for life?”. I gently share the truth with them. “No, they don’t; but they really have a strong allegiance to their territories!” The idea that loons love their homes, not their breeding partners, provides scant solace to most folks.

Having spent a quarter century disappointing loon lovers in Wisconsin and Minnesota, I have been searching for scientific news to share about breeding pairs that sheds a warm, wholesome light on loon mating behavior. My quest is not inspired by guilt alone. As a scientist, it makes sense to me that adults should become acquainted with their mates and benefit from doing so. How could complex, long-lived animals like loons jointly defend a territory, build a nest, divvy up incubation duties, and raise young together — as seen in Linda’s photo above — without benefitting in some way from their association?

At last, my current study of predictors of breeding success has revealed one way in which loon pairs do benefit from a long-term association. The graph below shows two patterns. First, there is a gradual improvement in hatching success over time as a pair remains together on a territory. These numbers jump around a lot owing to limited samples of pairs that have been together ten or more years.

Second and more clear cut is the improvement in hatching success between a pair’s first year together (Year 0) and their second (Year 1). As you can see, loon pairs improve their chances of hatching eggs by almost 10% between these first two years.

Now we can only speculate about the cause of this dramatic improvement in breeding ability among young pairs. Perhaps pair members synchronize their breeding behavior better in their second year together than in their first. Maybe pair members rotate incubation duties more crisply in year two, thus seldom leaving the eggs uncovered and unguarded.

Of course, the challenge that romantic couples face of living and working harmoniously, following an awkward adjustment period, has a familiar ring to it. That challenge is depicted in “Period of Adjustment“, a 1962 comedy-drama film based on a play by Tennessee Williams. It is heart-warming to see that loons — like actors Jane Fonda, Jim Hutton, Lois Nettleton, and Tony Franciosa — bounce back strongly after a rough first year.

I have had a lot to say about male loons and their experience. Indeed, the fact that the male decides where to place the nest means that he develops a tight bond to his familiar territory and fights hard — harder than his mate — to keep it.

But females too play a vital role in breeding success (like the pictured female photographed by Linda Grenzer on Bear Lake). How might female experience affect the outcome of a nesting attempt? Now that I have begun a detailed analysis of causes of breeding success and failure, I have started to ferret out the difference that female experience makes. I am only halfway done, but I can already see that the number of years a female has spent on her territory strongly affects the date her chicks hatch. As the graph below shows, females that have just arrived on a new territory — because they have evicted the previous female owner or replaced a dead one — have an average hatching

date of 22 June. In contrast, 4-year veterans on territories hatch their eggs, on average, 5 days earlier — June 17th. Now this might not sound like much of a difference in hatching date. But when you are tasked with stuffing your voracious chicks with fish, watching them grow rapidly to adult size, and hoping they get proficient enough at foraging, flying, and avoiding trouble to eke out a successful migratory flight to Florida, you take every extra day you can get!

You might ask: “Does this delay occur because of female inexperience with breeding in general or does it come about because of lack of experience on a specific territory?” The delay seems to be associated with lack of familiarity with a specific territory, because females that pick up stakes and move to new territories show delays in hatching date just like the ones they suffered on their first territories.

The cause of the improvement in nesting schedule with experience is likely to be energetic. That is, a female that knows how and where to find food on a lake is able to recover from spring migration quickly and begin the lengthy and arduous process of raising chicks. A female that is still learning where to find food on her new lake spends extra time before she reaches a suitable body condition to commence breeding.

Don’t male loons have just as big a problem restoring their body condition and thus readying themselves for a nesting attempt? Perhaps. But males experience minimal breeding costs until they begin joint incubation duties with females. Apparently males’ energetic deficit from migration does not hinder the breeding schedule.

You probably noticed that the graph depicts a curve, not a line. Older age-classes of females — those that have spent 5 or more years on a territory — actually begin to nest later than females that have spent 4 or fewer years there. What on Earth could explain this peculiar pattern?

Since it makes little sense that a female’s experience on a territory could begin to work against her after several years, we must look beyond experience for an explanation. The later hatching dates of more experienced females probably arise from reproductive senescence — a decline in reproductive performance that occurs with advancing age. Senescence is well-known in mammals and also many birds. We should not be surprised to see such a pattern in loons.

It is exciting to discover and ponder the reproductive quirks of female loons. Like many of our findings, this one only became visible because we studied thousands of nests, by hundreds of marked loons, across decades of their breeding lives. That, of course, is our bread and butter.

One of the quirks of the loon breeding system is that males choose the nest location. I have mentioned this fact many times in posts. I still scratch my head over the pattern, which makes no sense on many levels. Why would a breeding system evolve in which one sex or the other has sole discretion for deciding where the nest goes? In such a case, death or eviction of the choosing partner is devastating for its mate.

Consider this scenario. A male and female settle on a new territory on a protected cove in a large lake. At first they struggle to hatch young, but they begin to do so after a few years and enter a period of steady chick production. After 12 years, the male dies. Since he is the repository of information about safe and dangerous breeding sites in the territory, the veteran male’s death leaves his mate in the lurch. She must wait patiently while a new male takes over as breeder and stumbles badly for a few years by placing nests in the path of hungry raccoons. Ultimately, the new male learns by trial and error where eggs will be safe from predators, begins to reuse those favorable locations, and becomes an accomplished breeder. But, having endured several years of her mate’s reproductive ineptitude, the female has lost precious breeding time.

This scenario plays out incessantly in common loons. We field observers experience some of the frustration that veteran females must feel as we watch the poor nesting choices of their novice male partners. Until a few days ago, however, we did not appreciate the stark contrast between male and female breeding experience.

As the graphs above show, males do not simply improve in hatching success in their first year or two on a territory; they improve over a period of at least 20 years! And the cumulative impact of this steady improvement is massive. A male’s odds of a successful hatch are 35% better in year 20 than in year 1.

On the other hand, females do not experience better hatching success in year 20 on a territory than in year 1. In fact, there seems to be the faintest whiff of improvement for females over the first five years on a territory — and a tailing off around 20 years on — but these apparent patterns are not borne out by statistical tests.

By the way, these findings do not mean that a veteran female might not happen to enjoy higher hatching success as her tenure on a territory increases. What it means — and this can be confusing to think about — is that any boost in hatching success an individual female enjoys results from the growing nesting experience of her mate, not herself.

I am still scratching my head over this dramatic difference in breeding behavior between male and female loons. It still baffles me on many levels. But the dramatic, long-term gain in nesting ability of males does help me understand the viciousness of male territorial behavior. A male that has reaped enormous benefits while spending two long decades to learn the ins and outs of his breeding territory should fight desperately to keep it.

Sometimes during a night of capture, when we have finished color-banding a loon and are releasing it back into its territory with its family members, I say to the bird, half jokingly, “Welcome to the Loon Project”. But I mean it. Once we place colored leg bands on a loon, we start to feel a kinship with that loon and take an active interest in its well-being.

The bond we feel with each banded loon grows as team members report its trials and tribulations across many years of its life. “Red over blue-stripe on Lumen is soooo tame!”. “Omigosh, that female on Lumen was super aggressive when two intruders landed in her territory this morning”. “Red over blue-stripe really scared a kayaker that came too close to its chicks today”. “Red over blue-stripe fed its chicks 58 times during the hour I was observing the family; those chicks begged relentlessly.” “Red over blue-stripe just skulked around the southern end of the lake this morning while her mate foraged with a new unbanded female. She looked so bummed out.” “There is a new breeding female on Birch today; she is red over blue-stripe!”

Just as we mourn when a male or female is evicted from its territory by a young adult, we cheer when it bounces back and claims a new territory nearby with a new mate. If one of “our” loons should be injured by a lure or fishing line, we spring into action to save it.

Knowing and caring about our study animals makes it more enjoyable and rewarding to observe them. But the warmth and connection we feel towards our loons is really just a pleasant byproduct of a coldly pragmatic research philosophy: mark every loon you can, and track marked individuals obsessively throughout their lives.

Why are we so fixated on marking loons and studying marked individuals? Because marking and reobservation allows us to turn anecdote into science. If one watches five unmarked adult loons circling and diving together in early July on Brandy Lake, and two of the five birds yodel at each other, one might conclude that two members of the group must be males that became aggressive for some reason. If, on the other hand, the five loons are color-banded, we can begin to make inferences about behavior. We might observe that the group consists of two territorial pair members from Brandy and three intruders: a 3-year-old male floater reared on Johnson Lake, a 7-year-old male floater raised on Bullhead, and an 11-year-old female breeder from neighboring Arrowhead Lake. We might further note that the two yodelers are the 9-year-old territorial male and the Bullhead floater. And finally, we might observe that the 3-year-old and neighboring female fled from the group of 5 following the yodeling incident and flew off shortly afterwards, while the 7-year-old male engaged in many simultaneous dives with the male breeder and stayed 36 more minutes before departing from the lake.*

Of course, one visit to a breeding territory does not by itself lead to any useful scientific conclusions, even when loons are marked. But when this day’s observations are combined with those by scores of other field observers on hundreds of marked loons and thousands of early mornings, statistical patterns begin to emerge. Indeed, in a paper we just published, we document how floaters (nonbreeders too young to claim a territory) behave differently as they age, how territory owners tailor their aggressive behavior to floaters of different ages, and how loon parents optimize defense of chicks differently as they grow. So the accumulation of observations on marked, well-known loons made possible several steps forward in our understanding of territorial behavior.

Marked loon populations have value over and above the strides they help us make in understanding loon behavior. Since loon numbers have clearly declined in Wisconsin in recent years and apparently also among the less-well-known loons of Minnesota, our study animals in both states suddenly have special significance. In the coming years, we hope to use our study populations in Minnesota and Wisconsin to ascertain the causes of the declines and work with others who love loons to turn things around.

*Linda’s cool photo above is of Nelson Gould, a Chapman student, who worked with us for three years.

I have pointed out many times how science proceeds not in simple linear fashion but haphazardly. A finding may appear rock solid but subsequent findings modify our understanding, even forcing us to discard earlier conclusions in some cases. Viewed from space — and over large stretches of time — we might seem to advance steadily in understanding a phenomenon. From ground level, progress in comprehending a topic is herky-jerky; we make clear progress for awhile, then reach a dead end, back up, and find a new path forward.

So it has been with loon conservation. Thirty years ago, we feared that methylmercury was a major health hazard for breeding loons and their offspring. At the time, our fears seemed well grounded. Now, having spent millions of dollars measuring mercury levels in loons and looking for its impacts, we must reassess. We now see that, at worst, mercury has the potential to harm loons in the eastern 1/3 of the breeding range — and even there only in acidic lakes. Consequently, where loon populations appear to be declining, we can cross mercury off the list of threats and turn our attention to other potential causes. By the way, it was no mean feat to cross mercury off the list of dangers to loons. Thoughtful, rigorous work by dozens of biologists across North America has made this conclusion possible.

Our study of a much narrower topic — the loon yodel — also seems to be moving forward by fits and starts. (Linda’s gorgeous photo above shows a male yodelling in typical crouched position.) A really cool paper by Jay Mager — a collaborator with the Loon Project for several years — showed fifteen years ago that the frequency of the yodel was strongly dependent upon body mass. Large males, Jay found, had low-pitched yodels. This was fascinating to us, because it meant that male loons were revealing their size to territorial competitors! Now it is well and good to advertise your size if you are a big loon. In that case, the fact that you are yodeling tells opponents that you are motivated to defend your territory, and the low frequency of your yodel informs them that you can back up that determination with beef! But consider the mixed message that a small male sends by yodeling: “I am small but feisty”. Hmmmm. Not such a deterrent to a territorial challenger, it would seem. We are still coming to grips with this so-called “honest-signalling” of body size and trying to learn how small yodelers might benefit from yodeling.

Among males whose ages are known exactly (blue) and males whose ages are estimated (red), older birds have higher-pitched yodels.

But wait! Lately we have had cause to wonder whether body size really does have a strong impact on yodel pitch. Now that a good many males that we marked as chicks have settled to breed, we know the ages of many of our breeding males precisely. Brian Hoover, a postdoc here at Chapman, took these new age data — data unavailable to Jay Mager — and looked to see whether age is correlated with yodel frequency. As you can see from the figure, age is strongly correlated with yodel frequency: old males have high-pitched yodels. Reassuringly, the better our age information is, the stronger the correlation. That is, the blue points show a tighter age/frequency relationship than the red points.

Where does this leave us? I just explained that small males have high-pitched yodels. Now I am telling you that old males have high-pitched yodels. Can it be both? It is possible that both mass and age influence yodel frequency and could exert additive effects. (Imagine the extreme falsetto of a male that was both old and small!) However, Brian examined mass in his more complete analysis and found no evidence that it affects yodel frequency after all. In other words, we might have found a mass/frequency pattern back in 2006 simply because we did not have age data that would have shown us what was really an age/frequency pattern. Statisticians are quite familiar with this frustrating phenomenon; conclusions can change dramatically when a variable that was left out of a previous analysis (age, in this case) is added.

The finding that it might be age, not mass, that affects yodel pitch would be more palatable if high-pitched yodels by old males made more sense than high-pitched yodels by small males. But, intuitively, the new finding makes less sense! This is the delightful pickle we now find ourselves in. “Delightful” because the age pattern is robust and clear, as the graph shows. A “pickle” because we have no idea, at present, why or how old males have high-pitched yodels.

Well…so it goes in science!

I try not to steal a glance through the lab window each time I pass. But I usually fail. You see, Marco Bisoffi, a molecular biologist and colleague of mine at Chapman, has restarted our study of telomeres* in loons as a possible tool to measure age and the effect of stress. Each week Marco churns out telomere measurements on a new set of loons, as he tries to troubleshoot the PCR** procedure. So when I walk by his lab and see him bent over his laptop, I wonder whether his promising early finding that telomeres indicate age in loons has held up.

It has. Now that Marco has run twelve males and ten females of known age, the trend is stronger than before. If you study the plot above, in fact, two patterns are evident. First, old males and females have shorter telomeres than young males and females. Second, males as a group have shorter telomeres than females. (This latter finding repeats what Jeremy Spool had found a few years ago.) There is some scatter in the data, especially among females, but both patterns show high statistical significance. Of course, we will have an even better fix on these patterns when we have run the other 83 DNA samples we have collected from adults of known age.

It is hard to exaggerate the value of these findings for loon biology and our own research in Wisconsin and Minnesota. There are countless benefits to studying loons, but one drawback has always been our inability to “age” individuals effectively. To our enormous frustration, we cannot even distinguish a 5-year-old from a 30-year-old. If this telomere pattern holds up, however, that source of vexation will be considerably diminished. In the future, we will be able to take a DNA sample from an unknown adult, measure its telomeres, and assign it to an age-class. Indeed, if the unknown bird is a male and we record both its yodel and its tendency to yodel at intruders, we shall be able to narrow its estimated age range still further — probably to within a few years.

Why does it matter that we are on the brink of being able to age adult loons accurately? First, age has a strong effect on a great range of behaviors, including aggressiveness, ability to hold a territory — which increases in young loons and then declines later in life — and even willingness to incubate eggs when black flies are abundant. Second, age impacts survival rate, especially in males. So knowing the ages of loons helps us refine our estimates of survival and improves our models of population dynamics.

Speaking of age and decline, the featured photo for this post is Linda’s Grenzer’s pic of “Clune”, the male on her lake. Despite the inevitable shortening of his telomeres, this 23-year-old still looks pretty fit in his winter attire!


*telomeres — protective DNA sequences (“end caps”) on chromosomes that permit DNA to be replicated many times but become shorter with age and stress

**polymerase chain reaction — a common molecular technique that permits efficient study of specific regions of DNA

He was the biggest, healthiest juvenile we caught in Minnesota last year. The Rush Lake-Northeast chick was so independent on July 16th, when we first attempted to catch the family, that we could not relocate him after capturing and banding his parents. We shrugged, returned the following night, and had better luck. At 2900 grams, “Copper-White”, as he became after banding, was 300 grams heavier than the second-heaviest chick we caught last summer and almost certainly a male.

Considering the risky environment inhabited by juvenile loons, it is a mistake, I have found, to become attached to them. So, with the exception of the “Miracle Chick” — a juvenile on Squash Lake in 2012 that lost his father at three weeks, watched his mother quickly re-pair with a new male, but still got enough food and attention to fledge — we have tried to avoid this practice. Still, Copper-White became lodged in my mind. I had great hopes for him. If any juvenile had a chance to fledge, migrate, and come back in a few years as an adult, it was Copper-White.

Large size and good body condition, it seems, are not enough to protect a loon in his first few months of life. Last Friday, the National Loon Center got a report of a loon hemmed in by ice on on Cross Lake. They raced out to check the bird, and Mike Pluimer snapped the photo above.

It was alarming enough to hear of a loon still on the breeding grounds in mid-December. By this time, loons from the Minnesota population should have arrived in Florida and begun adjusting to a saltwater diet. Our hearts sank a bit further to see the bird’s plight. Resting in a tiny pool of open water surrounded by encroaching ice, this juvenile was clearly in dire straits. Why had he failed to migrate south with others of his species? Something must have gone horribly wrong.

Following heroic efforts on the part of the Crosslake Fire Department, Copper-White was caught and transported to Wild and Free Rehab Center in Garrison. Terri and Richard, who live on Rush Lake and watched the chick grow from its earliest days, reported that the captured bird was strangely docile — another worrisome sign.

Arrow points out where Copper-White’s right wing was sheared off at the metacarpal bone by a boat propeller. (Photo courtesy of Wild and Free Rehab, Garrison, MN.)

It took little time for Katie, the vet at Wild and Free, to diagnose Copper-White’s problem. The end of the loon’s right wing had been sliced off some time ago by a boat propeller, rendering him incapable of flight. Unlike many hawks and owls, loons’ size and need for open water make them impossible to keep alive in captivity. The only option was to euthanize this bird.

Alas, I have no cheerful anecdote to cushion the blow. We are disheartened to lose a healthy, strapping juvenile loon to a boat strike. But boat strikes that injure loons are a fact of life in the Upper Midwest. We lost a healthy adult male even more tragically two years ago in Wisconsin. The only comfort here is that boat strikes occur infrequently enough in the Upper Midwest that they do not contribute meaningfully to loon mortality. At the moment, that is cold comfort.

Although it has been over a decade, I still remember that morning vividly. I was observing the banded male and unbanded female on Brown Lake as they foraged on the wide portion of the lake’s eastern side. As is the case with most of our study animals, the loons were quite tame. They reacted indifferently to my red canoe as I tracked their progress slowly down the lake.

The loon pair’s relaxed foraging seemed odd during what had been a most tumultuous year on Brown. Though the female had resided on the lake since April, three different males had vied for and held the position of male breeder for portions of the season. Ultimately, “Mint-burgundy over Silver, Green over Blue-stripe” (Mb/S,G/Bs for short) drove off his competitors and became the male breeder. Evicted from Two Sisters-West in 2008, Mb/S,G/Bs had drifted about for two years before finally seizing control on Brown. Sadly, his victory in late June 2011 came too late for successful nesting to occur. So on the day of my visit, August 3rd, Mb/S,G/Bs and his mate were merely killing time before molting and readying themselves for the southward migration.

As I watched the laid back pair forage, an intruder appeared overhead. The pair watched the intruder as it slowed, descended, and parted the water’s surface to land twenty meters away. The intruder — a female hatched and reared 15 km north on Moon Lake, near St. Germain, three years earlier — was clearly uneasy. She bowed her head, dipped her bill in the water repeatedly as she drew near the pair, and initiated many brief dives as she circled them. For their part, the male and female breeder seemed to be going through the motions. They circled slowly with the intruder and peered at her when she dove but seldom dove themselves. The video below from South Two Lake depicts a similar scenario.

Afterwards I reflected upon the encounter. More clearly than ever before it seemed to me that I was watching a jittery youngster confronting two old, confident territorial loons. I am not sure why it had taken me eighteen years to do so, but I felt that I suddenly understood something very fundamental about loon territorial behavior. Loon pairs watch the behavior of an approaching intruder closely, quickly size it up — estimating the level of threat it poses to their territorial ownership — and then behave accordingly. As a result of this particular lake visit to Brown, my research team began to recognize and record “initiates dive” behavior (i.e. being the first loon in a group to make a short dive) and also “declines dive” behavior (refusing to dive when another loon nearby has done so). These advances led to new data collection and new insights into age-related territorial behavior.

Intruders, we have learned recently, provide ample signals of their age, fighting ability, and level of interest in battling for territory ownership. As the above figure shows, one of the clearest hallmarks of youth among intruders is the “initiates dive“ behavior. Young, timid intruders with no intention of vying for territory ownership are nervous Nellies, like this three year-old female was, and carry out many initiates dives. At the same time, these youngsters almost never show the “simultaneous dive“ behavior (which signals a willingness to escalate conflict), nor do they yodel or show aggression of any kind. Without question, there are dozens of other small signals that territorial pairs pick up from intruders to assess their age and degree of threat they pose.

And territory owners act upon the information they glean from intruders. That is, they treat a harmless visitor within indifference; they behave aggressively toward a dangerous intruder. The keen ability of territory owners to distinguish between intruders helps us understand how they can survive hundreds of intrusions each year without becoming exhausted. They save energy where they can and only get worked up and aggressive when they must.

These conclusions might sound obvious and intuitive. They are. And yet it took some 20 years and dozens of statistical tests to recognize and analyze the age-related patterns in behavior that allowed us to infer how both intruders and pair members betray their motives and strategies during such encounters. Fortunately, our perseverance has been rewarded. A few days ago our paper on interactions between intruders and territory owners was accepted for publication in a good behavioral journal. It should appear in print early next year. Thus, we are incrementally closer to understanding the entire territorial system of common loons.

If you would like to support our work in understanding territorial behavior, measuring population parameters, and conserving loons in the upper Midwest, consider a donation to the Loon Project HERE. At the moment, we are hoping to buy two canoes and a small motorboat, which would allow us to continue our long-term Wisconsin research while adding new lake coverage in our new Minnesota study area in 2022. Thanks for any support you can give us!