We all focus most of our attention on breeding pairs and their chicks. Why not? Breeders are the loons we get to know — day after day, year after year — as they struggle to hold their territories, choose nest sites safe from raccoons, incubate their eggs for 4 long weeks, and then protect helpless offspring from all manner of environmental threats. We admire the toil and turmoil they face each year and are downcast when they lose their breeding position, nest, or chick. And we grieve when they die. I am probably unusual in this regard, but I am also inspired by observing the challenges that breeding pairs routinely confront and overcome.

In our obsession with breeding individuals, though, we forget about the many loons living on the margins. These are “floaters” — mostly 2 to 7-year olds who spend the entire summer without a fixed home. Floaters are the individuals that forage alone on small lakes or skulk along the outskirts of defended territories, occasionally socializing with or accosting territory holders. Unlike territorial loons, they drift about.

Although their lives might seem simpler and less stressful than those of territorial loons, floaters — even young ones — face challenges of their own. Our work has shown that 2- to 4-year olds are much lighter than 5- to 7-year olds, are more submissive to territory holders during territorial intrusions, and almost never initiate battles for territorial ownership. Yet these youngsters do intrude into breeding territories. We presume, therefore, that even as they mature, young floaters collect information about owners. Our data show that young floaters intrude strategically into territories within a focused area (usually about 10km in diameter, see figure below from our recent paper) so as to meet and interact with owners of their own sex that they might

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evict down the road.

Having reached optimal adult condition, floaters of 5+ years of age begin to size up owners with greater urgency, choosing to battle those that appear weak or are noticeably weaker than they were on a previous visit. Male floaters probably also listen to the quality of a male owner’s yodel, because certain acoustic elements of the yodel convey information about the yodeller’s body size and condition. As we have seen repeatedly, intrusions by floaters of 5+ years of age are not welcomed by territory holders — and can be dangerous for both parties.

Considering that 2 to 4 year-olds are still reaching optimal body condition and 5 to 7 year olds are putting themselves at risk by actively seeking territories, it is surprising that floaters of both age groups survive at a rate just as high (about 90% annually) as established pair members (see figure below). Apparently the risks of probing and competing for a territory among young loons are roughly equivalent to those that come with


territory defense and chick-rearing among older individuals. (Note from the figure that only older age-classes, and only males, show lower survival.)

We are used to the invisibility of floaters. That is, we see and study them as intruders into defended territories, but we seldom consider where they come from or how they live. So I always get a jolt when a dead floater turns up, like the tame 5 year-old male in Linda’s photo from the Lake Nokomis area. These rare unpleasant finds are a good reminder that gaining a territory is a long and difficult slog.


Since snow and ice lingered far too long this spring, loons were late to nest in northern Wisconsin. The lateness of the season has also reduced opportunities to renest after early nest failures. There was simply limited time for pairs to weather four weeks of incubation and still rear the chicks to a point where they could learn to fend for themselves and make their way to Florida. Add to the narrow window this year the mishaps that cost us young chicks on several territories….and I was beginning to sweat.

But my fears of another off-year for nesting success in the Upper Midwest loon population have not been realized. A relatively short period of black fly abundance has helped immensely. As our recent paper showed, abandonments from black fly harassment are a good predictor of nesting success; that pattern has continued in 2018. So it seems likely that this year will break our four-year breeding slump.

Here are the numbers. As of July 4th, we had an estimated 48 breeding pairs in our study area with a chick or chicks. Eighteen (give or take a few) were still on nests, of which over half will produce chicks also. That leaves us with roughly 60 loon pairs with chicks. A handful of the 60 pairs will lose their chicks before 8 weeks; in addition, though, we will make roving visits to non-study lakes within and just outside the study area and find about 6-8 more pairs with chicks. When the dust has settled, we should end up with a number of pairs with chicks that is very close to the 65 successful breeding pairs we were able to band in 2013. That fact is worthy of note, because 2013 was both a year in which our procedures and lake coverage were similar to this year, and the last solid year of breeding. So we can all breathe a great sigh of relief — and enjoy Linda’s cool photo of the freshly-hatched chick on Muskellunge Lake and the female, with moist eggshell fragments still adhering to her breast.

As of today, 32 of our territorial pairs have hatched chicks. We are on pace with last year, despite the three-week delay in nesting resulting from the cold winter. I am encouraged; 2018 looks to be at least an average year for chick production.

My optimism about the breeding season has been tempered of late, as we have gotten fresh evidence of the frailty of newly-hatched chicks. In the past two weeks, four pairs with tiny chicks have lost them — a higher than average rate of loss. Linda first reported that the chick on her lake suddenly disappeared; her observations suggest that a snapping turtle might be the culprit. Yesterday, the chick on Crystal Lake (Lincoln County) fell victim to the attack of an intruding loon, as reported by a lake resident. And two northerly pairs — on Blue-Southeast and Carrol — also completed four weeks of incubation only to lose their chicks in the first few days after hatching. The causes of chick loss are unknown in these two cases.

Loss of chicks in the first two weeks of life is not terribly surprising. Young chicks must confront a great many challenges, including simply keeping themselves warm and avoiding physical injury as they learn to swim and move about. But the greatest hazards to hatchlings, we have learned, are strictly biological. Being tiny, having limited mobility, and with only a vague sense of the dangers posed by much larger organisms in their habitat, young chicks can be attacked and killed by a wide variety of animals — intruding loons, snapping turtles, muskies, and eagles, to name a few. In contrast, adult loons are wary, agile, and large enough that the list of potential attackers is short. Adults have simply outgrown much of the danger from other living things in their environment.

As discouraging as it is, I am convinced that this recent spate of chick deaths is a statistical blip. There has been no sudden change in the loons’ habitat, or explosion in the population of any predator, that could signal the beginning of a worrisome trend. Rather, this is just an unfortunate negative swing that will be washed out by later swings, negative and positive, and ultimately forgotten. Perhaps we can hurry the process along by focusing on Linda’s recent photo of the female (“Honey”) and chick on Muskellunge Lake.

What is better than finding out that your just-published article has been featured by an online science media outlet? Finding out from your dean! An hour ago this happened to me as I strolled out of our new science building.

We were excited to learn this article has excited some attention. It was a bit of a sleeper. Published in a good — but not spectacular — journal, our investigation of the flies’ impacts and loons’ logical responses to them caught the eye of the journal’s media department. I will not bore you by rehashing our findings, which I have discussed before. By the way, a related media blurb included Linda Grenzer’s cool photo of the male on her lake sticking on the nest in 2017 despite flies biting him mercilessly. (Another of Linda’s related photos appears above.)

Fortunately, this year has been a mild one for black flies. So while pondering the harsh negative impacts that black flies often have on loon nesting behavior and breeding success, we can all relish their absence.


Most of us have been there: determined to take the next step in life but thwarted in our efforts to do so. More often than not, another individual occupies the position we seek. That is the situation in which nine year-old W/G,B/S (hereafter “White-Green”), finds herself on Blue Lake.

Blue seems a good fit for White-Green. She grew up on Franklin Lake in Forest County, another large, clear-water lake 35 miles to the northeast. Since young males and females try to settle on breeding lakes similar to their natal ones, it is natural that White-Green should try to find a place on Blue.

But setting your sights on a goal and achieving it are two different things. Back in 2014, when she was only five years old, White-Green battled with the 15 year-old female from the Blue-West territory and lost. Since then, she has made sporadic attempts to evict her decade-older rival, failing each time. In 2016, she observed the violent overthrow of the male at Blue-Southeast by a younger rival. However, the carnage that ensued offered White-Green no opening at a breeding position at Blue-Southeast, although she paired briefly with the male evicted from that territory. Since then, we have so often seen White-Green on Blue Lake – and never on adjacent lakes that we also monitor — that she must live there, hiding out in the swath of unoccupied space between the two breeding pairs. And waiting.

She waits still. Two weeks ago, White-Green again challenged the Blue-West female, in what has become an annual ritual. I have begun to feel frustrated on White-Green’s behalf. Her many fruitless attempts at usurping a space recall times when I myself have been baffled in a crucial life pursuit. It would seem that the writing is on the wall: White-Green is not strong enough to subdue the breeding female at either end of the lake and seize their breeding position. Shouldn’t she move on? Isn’t she wasting valuable time in the prime of her life? As a biologist, I come at this from another angle, asking, “Could White-Green’s obsession with acquiring a territory on Blue Lake be evolutionarily advantageous?”

A precise answer to this question would require a calculation of evolutionary fitness benefits of fighting for a known resource versus turning her attention to a new target — a neighboring territory on Bobcat, Bolger, or Kawaguesaga. We do not have sufficient data to make such a quantitative comparison. However, White-Green’s “siege” of the Blue-West territory does highlight two quirky features of the loon breeding system that might help us understand her odd behavior.

Whether male or female, nonbreeding loons in search of a territory set their sights on a handful of them that they visit often, rather than casting a wide net that includes dozens of potential territories. We infer from the narrow scope of their search that young nonbreeders might use a strategy of sizing up territory owners, monitoring their condition regularly, and challenging them for ownership when they seem most vulnerable. (Of course, nonbreeders would prefer the low stress alternative of replacing a breeder that has died.) The tradeoffs of focusing on a small number of territories are clear. By getting to know the behavior of a few breeders well, young nonbreeders might be able to pick up subtle changes in breeder behavior that signal weakness and allow them to time their eviction attempts effectively. And conducting a narrow search might also permit a nonbreeder to learn about the illness or death of a territory owner rapidly, so that they can quickly mount a bid for the territory and claim it before others learn of the opportunity.

Another quirk of the loon breeding system is a female-biased sex ratio caused by greater longevity among females. That is, there are always more breeding females than breeding males in the population, since many males die young. The longer lives of females is a double-edged sword from their standpoint. On the one hand, there is vigorous competition for any female breeding vacancy that becomes available. On the other hand, females live long enough that what at first appears to be an unhealthy obsession with one territory or two might ultimately be rewarded.

Breeding prospects for loons in northern Wisconsin seemed dim only three weeks ago. Not only had a frigid April delayed the start of nesting, but Simulium annulus was doing its best to keep loons from warming the eggs that had taken so long to appear. A statistical correlation between cold spring temperatures and black fly harassment had me fearing that the long-awaited nests would be abandoned in short order – delaying the season still further. My hopes for a bounce-back year of breeding, after 2017’s disappointment, seemed distant.

As I keep learning in life, unfathomably horrid situations often improve. So it was this spring. To be sure, loons were forced to abandon a few early nests – those at Langley, Fox, and Wind Pudding-East, for example – owing to fly harassment. But loon pairs that had been reluctant incubators in mid-May suddenly bent to the task late in the month. Even after accounting for the inevitable wolfing down of eggs in exposed nests – such as those at Two Sisters-Far East, Long, and Little Bearskin — by raccoons and their ilk, the vast majority of our breeders are sitting on eggs (like the male on Linda’s lake; see photo). At last count, 79 of 123 pairs we cover are on nests that have survived the crucial first ten days. Two weeks or more of incubation remain for most of these territories. But barring some unforeseen disaster, 2018 might be one of the most productive years for northern Wisconsin loons in the last quarter century. Who would have guessed that a breeding season that started so inauspiciously would gain such momentum?

The Nose Lake male I observed today had a conspicuous scar on his head. This particular scar, which I dutifully sketched on my datasheet, was located on the right side of his head, behind and beneath the eye. In comparison, the head of the Nose Lake female was sleek and without blemish. Scarred males paired with pristinely-plumaged females are a common sight. In fact, the scar I recorded today was the 72nd of the study – and the 63rd seen on a male. Moreover, this scar has persisted for a month; Linda photographed this bird on May 6th, and the scar was obvious then.

Scars on heads of males, which occur when they grasp each others’ heads and necks in a territorial battle, bring to mind the short, violent lives that many males lead. Slowly and surely, we are beginning to understand why male battles are fiercer than female battles. Part of the explanation for this pattern has to do with nesting behavior. We know from analysis of nesting behavior among color-marked breeding pairs that male loons control the placement of the nest. While we do not know why males control nestsite placement, we can see that male control of nest placement cranks up the stakes for male territorial battles. Why? Because male loons learn by trial and error where to place the nest. Once a male has nested successfully on a territory, he reuses that good nest location again and again, boosting his hatching success. Therefore, once established on a territory where he has nested successfully, a male has a large stake in holding that familiar territory. If evicted from there, the male must relearn where to nest and where not to nest on a new territory, which costs him precious time and energy. In contrast, females, which do not control nest placement, can freely move from one territory to another without paying a penalty in lost familiarity and, hence, breeding success. Since one territory is, in effect, as good as another to them, females should fight less hard than males to remain on a territory – and they behave as predicted.

A second part of the explanation for violent male battles is rapid senescence. Again, while we do not yet understand why males should age so badly, compared to females, the contrast in senescence has strong implications for male behavior. A male reaches a point – in his mid-teens typically —  where he is in rapid decline. That is, he is losing body condition and is at great risk for losing his territory. With the future offering little reproductive promise, many males in their mid-teens increase their aggressiveness and territory defense so that they can squeeze another year or two of breeding out of their territory. This, of course, is the terminal investment finding that I have been blogging about for the past months. (By the way, that paper has just been published online.)

With two factors – male nestsite selection and senescence – at play, we can begin to understand why males might be so violent. The factors are additive. A fifteen year-old male on a familiar territory is both falling into decline and facing a steep loss in breeding success, if evicted. So he has two good reasons to fight like hell to hang on.



Last week was a tense one. Dozens of pairs had laid one or two eggs, but black flies descended on them, making incubation impossible for most. Of the fifteen or so pairs with nests last week, only two incubated at all, and one of these pairs sat only during the first twenty minutes of our observations. (The other was Linda Grenzer’s notorious overachievers, Clune and Honey.) All other pairs spent their time in the general vicinity of the nest — but diving frantically, shaking and tossing their heads, even snapping their bills fruitlessly at the relentless biting insects.

Already frazzled by the grading of 131 all-essay final exams, calculation of final grades, and strident pleas of disappointed students — I fielded another while writing this post — I feared another dreadful year of nest abandonments, like 2017.

What a difference a week makes! While not altogether gone, the flies are dwindling rapidly. Breeders that could only view their eggs from afar 7 days ago are back on them. I caught the male in the photo — from Two Sisters Lake-Far East Territory — about to hop cheerfully onto his nest, after an incubation switch. He and his unbanded mate still have 27 days of incubation remaining. But, like all of our focal pairs, they have overcome the first major hurdle.

Spring is on its way. As I twiddled with my phone just now to check conditions in the study area, the temperature was nine degrees warmer in northern Wisconsin than in southern California — the result of an unseasonably warm day in the study area and an unseasonably chilly one here in SoCal. At long last, the ice is melting, and loons will soon be back on their lakes. Meanwhile, as Linda’s photo shows, they gather in droves on open water. Being Linda, she also checked out their bands, yesterday and today, to see how many she could identify. We are all the beneficiaries of her tireless efforts, because her IDs provide a window onto the lives of returning breeders that are, for the moment, baffled in their efforts to reclaim their territories.

Who are these birds, and where are their territories? Are they males or females? Are they, in fact, breeders or young floaters, who lack territories and will spend the year challenging territorial residents for ownership? Are they migrants hundreds of miles from their breeding area or neighbors stopping by until their nearby lake becomes ice-free? We cannot answer all of these questions, but we can answer most of them based on Linda’s meticulous observations.

First of all, the loons in these aggregations are almost all males. Of the 12 birds positively ID’d by Linda, 10 are known males, one is a female, and one is of unknown sex. Linda suspected this herself: she reported numerous territorial yodels, sometimes by multiple males at once, as the birds rafted about in ice-free portions of the lake. Females must be only a few days behind; in fact, as I was writing this, Linda reported seeing a breeding pair near last year’s territory on Nokomis.

Second, loons present on the breeding grounds at the cusp of the seasons are mostly breeders that have come to reoccupy territories, rather than young floaters bent on evicting them. Only one ID’d loon from Linda’s list is a possible floater; all others are known breeders.

Third, loons that aggregate on ice-free water in early spring are a geographical smattering — many from neighboring lakes but a good many from farther north, whose lakes might not be open for a week or more. Linda’s sightings show this clearly, as her list includes breeding males from:

1) the adjacent Nokomis-East Central territory,

2) Indian Lake, which is 6 miles to the NE,

3) Soo Lake, 11 miles to the NE,

4) Silver Lake (Lincoln County), 8 miles S,

5) South Blue Lake, 16 miles to the N,

6) Miller Lake, 18 miles N,

7) Blue Lake-West Territory, 20 miles N,

8) Forest Lake (Vilas County), 40 miles NE,

9) Rock Lake (Vilas County), 50 miles N, and

10) Crab Lake (Vilas County), 50 miles N.

and a lone female from Burrows Lake, 10 miles NW.

What can we learn from these sightings? The preponderance of males confirms that males precede females on migration by at least a few days, and shows that males are the main ones bottlenecking on rivers and open lakes near their ice-locked breeding territories. Returning males push hard to return very close to the date that their lake becomes ice-free; females dawdle by a few days. Why it is so crucial for males to return as close to ice-out as possible is unclear, especially since the paucity of young, non-territorial birds present suggests that breeders are not likely to be challenged for territorial ownership in the first few days after settlement. Perhaps males, because they select the nesting site, return as early as possible to take note of any changes in the lake or shoreline over the winter that might require them to move the nest from the prior year’s site.

We can also discern that these early spring aggregations do not comprise neighbors reacquainting themselves after a winter apart, but individuals from far-flung lakes that do not know each other at all and are likely not to encounter each other again. While we should not be surprised by some tense moments between such unfamiliar individuals — it is, after all, spring and hormone levels are high — we should expect mostly peaceful loafing and feeding. These males simply have nothing to gain from battling unfamiliar loons.

In short, the rafts of loons in Linda’s photo are playing a waiting game. Since territory settlement is a time during which owners must defend their lakes vigorously — even if they do get a brief hiatus after settlement, owing the late return of young challengers — we would expect these males to do next to nothing until their lakes open up. They should feed, rest, and expend as little energy as possible before the onerous task of breeding begins.