2016-07-31 01.32.29

Loon capture is a blur. We set out from our house at 8:45 pm, launch our small motorboat on the first lake, wait for nearly complete darkness, and catch any loon chicks and parents that we can net easily. By the time we have repeated the process four more times, we are rubbing our eyes, our weariness justified somehow by the presence of the sun lurking just below the horizon.

As an essentially negative person, what I often recall after a night of capture and banding are the physical demands of the process and my complete exhaustion. But there are dimensions of the work that are exciting and rewarding. Each loon is unique, and one never knows whether an individual will permit itself to be approached closely and netted or will be wary and elude us. So we experience many disappointments, but they are tempered by the occasional thrill of capturing an individual that, at first glance, appeared too skittish to catch.

The fruits of loon capture are obvious. By marking individuals and resighting them year after year, we learn about survival rates of adults and juveniles, territory fidelity, natal dispersal, and habitat preference. We glean a good deal of important information from these data. For example, survival rates of young and adults allow us to learn whether the  local population is increasing, decreasing, or remaining stable. And tracking of young loons from egg to first territory has revealed that loons develop strong preferences for breeding lakes that closely resemble their natal lake. Finally, capture is essential as a means to disentangle loons that have been run afoul of angler’s lines or lures.

This year’s capture exposed another distinctive pattern in loon ecology: the presence of ecological traps. An ecological trap is a breeding habitat that appears at first glance to be a good one but ends up being poor for reproduction. For example, a field might experience a burst of insect activity during early spring, enticing songbirds to settle there for breeding, but a crash in insect levels after eggs hatch might occur that suppresses the number of young birds produced. Two nights ago, we captured two chicks from two different lakes back to back. The first territory was a shallow 11-hectare portion of Wind Pudding Lake (my favorite lake name). The chick captured there was a five-week-old that weighed a scant 0.92 kg — less than half what we would expect from a chick of that age. Our daytime observations show that the chick’s parents are no slouches; they respond to its constant begging by making frequent dives and retrieving what food they can to feed it. Moreover, the chick itself dives often to forage. But this shallow lake, covered almost entirely by lily pads (which impair loon foraging), offers scant sustenance. I am afraid that the emaciated Wind Pudding chick will ultimately starve to death, as did the chick on nearby Liege Lake, another shallow lake choked by vegetation. Loon parents on small, acidic lakes struggle to rear even a single chick, whereas those on large lakes of neutral pH often raise two. This stark contrast was highlighted for us, as the lake we visited following Wind Pudding was 1373-hectare Lake Tomahawk. To be sure, loon parents on Tomahawk must steer their chicks through countless jet skis, water-skiers, anglers, and speed boats at all times of day. But vigilant parents are rewarded with abundant food for themselves and their chicks. The Tomahawk-Sunflower Bay chick held by Mina in the photo weighed 3.02 kg, yet it was only a few days older than the chick on Wind Pudding. Clearly the strapping youngster in the photo is heading for a healthy future and likely fledging.

Why on Earth would loons settle to breed on lakes that often provide too little food for their chicks? The answer might relate to the disconnect between nesting and foraging requirements. Alas, large lakes that contain many fish for loons often lack the islands, emergent marshy bays, and bogs that allow loons to avoid egg predators like raccoons. So loons looking to breed seem to be lured onto small, marshy lakes that yield successful hatches but doom their offspring to starvation.

Why would a breeding pair of loons — which has made an enormous investment in defending a territory, finding a nest site, incubating their eggs for almost a month, and then rearing their chicks — leave them alone? Such behavior seems reckless, almost dysfunctional. Yet loon pairs commonly “desert” their offspring for periods of hours, while they themselves visit neighboring lakes. During such times, of course, chicks are left to cope with all manner of predators and other dangers without parental assistance.

The leaving of chicks behind by parents does not occur willy-nilly. That is, not ALL chicks are deserted, only those that have reached at least four weeks of age. Some chicks, like those I watched on O’Day Lake this morning, are highly alert to their surroundings, like adults, and capable enough hunters that they are already providing over half of their own food. (Note the expert-looking dive by the chick in the video.) One can imagine that temporarily leaving behind chicks of this age does not carry huge risks. At six weeks of age, chicks can detect and flee from eagles, when necessary. So perhaps the risk of leaving your chicks to fraternize with neighboring adults is not great.

Still, temporary chick-desertion surely carries some risk. If the choices are remaining on your territory with your chicks versus departing to a nearby lake to visit with other adults, the first option is clearly the safer one. The only conclusion to draw from what appears to be rather reckless socializing by pairs with chicks is that they must gain something nontrivial from doing so to compensate for the small risk that their chicks will be lost during their absence.

Their are several possible explanations for parents’ trips to neighboring lakes. One hypothesis maintains that parents encounter and become familiar with neighbors whom they might have to battle later for a territory or with whom they might pair in the future. If so, recognizing and learning about other adults might provide for more effective fighting or breeding. A second idea is that parents leave their chicks in order to forage on lakes other than their own so as to maintain robust food levels for the chicks. Thirdly, temporary desertion of the chicks might constitute part of the weaning process; chicks often beg incessantly when with their parents, and perhaps chicks must be without parents to begin foraging effectively on their own in preparation for adulthood. A fourth explanation is that parents, which are conspicuous to other adults owing to their bright plumage, desert their breeding lakes in order to avoid giving away the fact that they have chicks. (Behaving this way might reduce the likelihood that a young nonbreeder could target the territory for eviction, since we know that intruders use chicks as an indication of a good territory.  ) If this explanation is correct, then parents are essentially trying to “decoy” intruders away from their own lake by visiting a neighboring territory nearby. The hypothesis is plausible, because: 1) intruders are strongly drawn to other adults in the water, and 2) intruders appear to find chicks only after seeing and approaching the chicks’ parents.

One final curious behavior seen in loons and chicks provides partial corroboration for the “decoy hypothesis”. When faced with an intruding loon flying over their lake, adults and their chicks dive and scatter. That is to say, adults and their chicks make every effort to hide from intruders by diving and spreading out in the lake, which complicates discovery. If loon parents are desperate to hide their chicks from intruders, what better way to do so than by flying off to other lakes near their own — and thus using their conspicuousness to draw intruders to those lakes — and leaving their cryptic chicks at home, where they will most likely escape notice from the air.

The decoy hypothesis is complex. Its crux is that loon parents are protecting their own future breeding prospects by taking a slight risk with their current chicks. In time, we will be able to determine if parents that practice short-term chick desertion enjoy longer territorial tenure. If so, this will be a stunning example of effective long-term planning in the animal kingdom!



With Mother’s and Father’s days still fresh in mind, maybe this is an apt occasion to relate a quirky instance of parenting in loons. The topic came up abruptly last week, when Doug Giles, a lake resident in British Columbia, sent these photos. At first, I could not even

_IMG0771 fathom what species of duckling was receiving this unanticipated caregiving. A frantic web search of ducks breeding in the vicinity revealed that this is a young Common Goldeneye. But how could such a duckling escape the watchful eye of its own mother and blunder into the path of a pair of loons?

The curious breeding behavior of goldeneye ducks provides a clue. Goldeneyes lay 7 to 10 eggs in tree cavities, and the female is the sole incubator. (Male goldeneyes leave the picture shortly after incubation begins and play no role in rearing offspring.) Goldeneye ducklings follow their mother away from the nest a few days after hatching. Unlike loons, goldeneye young feed themselves; their mother merely protects them and guides them to feeding areas where they are relatively safe from predators. The task of herding ten or so ducklings about creates logistic hurdles; ducklings commonly fall behind or lose track of their mother and siblings. When goldeneye hens converge in the process of guiding their respective broods, young intermingle, creating creches of dozens of ducklings of varying parentage. Finally, female goldeneyes, especially those with small broods, often abandon them — leaving the ducklings to fend for themselves or coalesce with broods where the female is still present.



In short, rearing of massive goldeneye broods seems haphazard and impersonal when compared with the parental behavior of common loons. If common loons are modern-day human parents in the era of small families, abundant participation awards, and kindergarten SAT prep; goldeneyes are the all-out reproducers of the pioneer days, never quite certain of how many young they have and where they all are! So we should not be surprised to encounter a misplaced or forgotten goldeneye duckling.

But why would loons adopt a young animal that differs markedly from a loon chick? First, it is important to understand that loons — like many animals — are hormonally primed to care for young. That is, young production is the crucial measure of evolutionary fitness, so we should expect all species to be committed to the rearing of young, if parental care is obligatory for survival in that species. Apparently, the drive to provide parental care is sufficiently strong that it can get misdirected at times. Among all ducks that loons might adopt, goldeneye are a good match, as they are diving ducks (not dabblers, like mallards) and subsist mainly on aquatic invertebrates and occasional small fish. Clearly this duckling is developing more of a taste for fish than is usual for its species, as Doug Giles’ first photo shows. The duckling’s willingness to accept food from another individual, moreover, shows impressive flexibility in behavior that we might not have expected.

While we can understand how the adoption of a goldeneye by loons is plausible, it is still a remarkable event. The likeliest outcome of a close brush between a small duckling and a loon pair is an attack; many ducklings are killed by loons each year. Perhaps this loon pair was primed for adoption by having lost their second chick shortly before they encountered the stray duckling.

One factor to which we definitely cannot attribute the successful adoption is relatedness between ducks and loons. Ornithologists have long known that ducks and geese are not closely related to loons. In fact, goldeneyes are cousins of turkeys and chickens, while loons are in the evolutionary lineage that includes herons, cormorants and penguins. The superficial similarity in appearance between loons and ducks results from evolutionary convergence, not common ancestry. In my view, the distant relationship between loons and goldeneyes is yet one more reason to marvel at the odd parent-offspring bond that has formed here — and hope that it leads to a good outcome.


Mina caught the tail end of the battle. By the time he arrived for his observations, the territorial male on Blue-Southeast was on his heels. No doubt shell-shocked from a beating he had received from an aggressive four-year-old male that had lurked in the neighborhood for two years and was making a play for territory ownership, the territorial male beat a hasty retreat, actually flying off of the lake with another intruder.

The strategic retreat of the territorial male left his mate in dire straits. Without another parent to engage intruders, the female alone had to defend the week-old chick from the aggressive onslaught of the four-year-old male. The situation was hopeless, the suspense only fleeting. The young male quickly discovered the chick and — in the grisliest moment we have observed while studying loons — snatched the chick out of the water and carried it for a time while pursuing the retreating female. When he dropped the lifeless youngster, it was over.

When the shock wears off, we will one burning question about this episode: “Why is Blue-Southeast the site of such frequent territorial controversy?” Can it be coincidence? One must never be hasty to rule out chance, but I think that is not the explanation here. Rather, evictions of the territorial male in 2015 and the same individual again this year resulted from his relatively poor fighting ability. Intruding loons are adept at sniffing out weakness in territorial residents. When a breeding male or female is unable to drive competitors forcefully off of the lake, intruders congregate, leading to further confusion and attempted evictions, in a disheartening positive feedback cycle. The Blue-Southeast male, while a capable parent, has often encountered intruding males that he could not drive off of the territory in years past. It is quite possible that his troubles in 2015 occasioned more territorial challenges in 2016 by the same set of challengers.

I am by nature a rather negative person, always quick to point out the downside of every cheery situation. But I see one bright spot here. If the usurper of the territory and killer of the chick becomes the territorial male, he would appear to be a strong and aggressive individual and one likely to defend the territory with greater success than his predecessor. This 2012-hatched product of nearby Bolger Lake would be a record-setter too, since he would be the youngest male ever documented to seize a territory by force. (Most young males, like very old males, settle on vacant territories rather than fighting their way onto occupied ones.) I wish I could offer more meaningful solace to Blue Lake residents, who will be dismayed at today’s turn of events. The best I can do is to suggest that the future for the Blue-Southeast pair is likely to be brighter than the recent past.









LMG_8014 Fall Molting Loon

Some months ago, I reported with excitement that loons that leave their first territory — usually because they are evicted from it by another loon — have a strong tendency to settle on a second territory with similar water clarity to the one they left. It was a striking finding and one that would have added a fascinating story to ecologists’ limited understanding of how animals choose where to settle once they reach adulthood. That pattern, which initially seemed a robust one, has turned out to be trivial.

Let me explain. When loons are forced to shift from one territory to another (or do so voluntarily), they typically settle on a second territory very close to their first one. Often they move to the lake next door or even settle on a different part of the same lake. What we thought we had found was that those “resettlers” carefully choose to settle on second lakes of similar clarity to their first lakes. But such a pattern is tricky to show conclusively. In order to do so, one has to determine how similar 2nd lakes would have been to 1st lakes if loons had settled at random and compare the actual pattern to random settlement.

The problem we ran into was spatial autocorrelation. Spatial autocorrelation simply means that within a large area, two points that are physically close to each other resemble each other more closely than two points that are farther apart. It turns out that lakes of similar clarity tend to be clustered. That is, if one lake is clear, then its neighboring lake is likely to be clear also. The same is true of murky lakes. So while we thought that loons were purposely choosing to settle on second lakes of similar water clarity to their first ones, they were merely settling on lakes near their first one which happened to be similar in clarity to their first because of proximity. In other words, we have no evidence that choice of lakes is non-random.

It took us a long time to discover the error. In fact, Mike Palmer and I had written up this finding and submitted it for publication before we realized — after running a new analysis requested by a reviewer — that spatial autocorrelation was solely responsible for our result.

Naturally, I am feeling a bit like the female on Muskellunge Lake, photographed recently by Linda Grenzer. A month or so ago, this bird and I were on top of the world — she because she and her mate had just fledged two chicks (a bumper crop for loons) and was in her showy black and white formal attire; I because I had an exciting finding that I could not wait to share with colleagues. Now the luster of the recent past is gone. The female’s chicks have dispersed to fend for themselves, she has lost her gaudy summer plumage, and she is about to resume survival mode during migration and winter. I have lost my thrilling discovery and will have to pick myself up, dust myself off, and attack some other scientific question.

Luckily, the Loon Project supplies an endless string of questions. One that tantalizes me at the moment: do chicks that fledge and leave their natal lakes to forage on other lakes nearby favor those that resemble their natal lakes? Such a result, if it occurs, would dovetail nicely with our finding that the first lake a loon ever settles on as an adult (at 4-7 years of age) tends to be strongly natal-like in size and pH. This result (he adds quickly) is not confounded by spatial autocorrelation, because first lakes tend to be far from natal lakes. At any rate, I shall let you know what I find.

When loon biologists measure reproduction, they often count chicks older than five or six weeks of age as having fledged. This is reasonable. Even though loon chicks remain dependent upon their parents for 11 weeks or more, almost all chick mortality occurs in the first few weeks of life. If a chick makes it to five weeks, it is almost certain to make it to adulthood.

Not so during this past week. As the unpleasant photo by Dan Pagel shows, one of the the Blue-Southeast chicks died quite violently several days ago after a sudden attack by an adult. Based on the photo and the swiftness of the death, we suspect that severe heart or liver damage caused the death. Since Dan was a few hundred meters away when the murdered Blue-SE chick IMG_3960vicious attack occurred, he was unable to identify the perpetrator. Regardless of what loon committed this act, it comes as a shock. The departed chick was over twelve weeks of age, healthy, and off by itself feeding when it fell victim to the brutal assault. Twelve weeks is much older — I believe as much as eight weeks older — than we had ever observed a chick to be killed by a foreign adult. In light of the tempestuous territorial scene at Blue-Southeast recently, it is tempting to view this chick as a casualty of the conflict that caused its father to lose his territory. Certainly, the new, unmarked male that had taken over on Blue-Southeast and had already been observed to peck the chicks in his first weeks of ownership is a suspect. A second possibility is that the chick carelessly strayed into the west territory on Blue Lake and was attacked by a member of the West pair for territorial reasons. But territory defense is scant and subdued at this time of year, so that explanation is problematic.

I was still coming to grips with the events at Blue Lake a few days ago when I received a second piece of news, equally discouraging. One of the chicks at Buck Lake, near Rhinelander, had died. The cause of death in this case is less clear; the chick’s leg became injured a few weeks ago — a lake resident reported that a snapping turtle latched onto it! — and the chick was not able to move about effectively afterwards. Like the Blue chick, the Buck chick was far past fledging age, having reached 11 weeks.

What can we conclude from the co-occurrence of two deaths in very large and healthy chicks? I think these two unfortunate events simply remind us that, while chicks that have reached near-adult size are far more likely than 2-3 week old chicks to survive to adulthood, they are still young animals. As we have reported in a scientific paper, loons suffer a higher rate of mortality during their early years. Adult loons have had years to hone their ability to feed themselves and cope with environmental hazards, including conspecific attacks; youngsters are still getting it down.

LMG_7851 Walter n Sammy

The season began inauspiciously at Crystal Lake in Lincoln County, which is along the southwestern fringe of our study area. Though it is a pleasant, rather clear lake, Crystal offers no nesting habitat for loons. Loons love most of all to place their nests on islands but also make use of boggy or marshy areas, which seem to provide safe havens from egg predators like raccoons and skunks. Crystal offers none of these options, as its shoreline comprises upland habitat and is thus easy for a hungry mammal to patrol. In fact, Crystal is such an inhospitable place for loons to breed that loon eggs vanish from its shoreline almost as soon as they are laid.

So I had mixed feelings to see a six year-old male hatched on nearby Muskellunge Lake settle to breed on Crystal in 2012. On the one hand, I was pleased that the young bird had found a territory after wandering around the local neighborhood of lakes for the three previous years. On the other hand, a settler on Crystal seemed doomed to reproductive failure. But the male and his parade of mates (three females in five years) persisted in their breeding efforts. As I reported earlier this summer, the male and his new mate in 2015 (she still licking her wounds from a recent eviction from neighboring Deer Lake) took a new approach to nesting in 2015 by choosing to place a nest on a swim raft off of a resident’s dock.

LMG_6672 dad n chick on raft

Once the egg had been protected from rolling off of the raft with a ring of rubber cut from an old tire and some natural nesting vegetation (thanks to quick thinking and skillful craftsmanship from Linda and Kevin Grenzer), things began looking up for the breeding pair. Although it was a bear for the male and female to clamber up onto the swim raft to incubate the egg, the setup seemed little different from the artificial nesting platforms that many lake residents place on the lake each year for their nesting pair to use. Given the virtually impregnable location, the probability of hatching a chick had gone from almost zero to near one.

As expected, the chick hatched successfully, and though it had trouble maneuvering around the rubber-reinforced nest (as Linda’s photo shows, above), it left the nest with its parents within a few days. Only when Linda reported that Crystal was experiencing a burst of territorial intruders — one of whom, by chance, was the mother of the Crystal male and hence the grandmother of the chick — did she and I become panicked. Intruders pose a grave hazard to small chicks. Would the pair that had so miraculously pulled off a hatch turn around and lose the chick to an infanticidal visitor? The threat was not trivial. As Linda’s remarkable and chilling photo shows, the Crystal parents broke a very basic rule of loon parenting: never permit intruders near your chick when it is less than 2 weeks old.

LMG_6709 Two intruders approach parent n chick2

Yet the chick survived this brush with death. Nourished by a healthy supply of fish from the lake, the chick grew so rapidly that we were able to band it at three weeks of age, much younger than is usually possible for loon chicks. It has continued to flourish since capture. Now, at roughly 8 weeks of age, it looks like a 10-week-old, as the opening photo attests.

I have a tendency to dwell on disasters. Confronted with a large set of events — some positive, some negative — the latter seem to stick with me longer, leaving me with a sense that things are not going well at present and might not go well in the future. Yet at the end of a season when many breeding loons flirted with disaster — and one died — the story of the plucky Crystal chick stands as an odd and memorable exception.

If you are following the events on Blue-Southeast, you will recall that the male there, who was caring for two large, healthy, 5-week-old chicks, was evicted and seriously injured by an unbanded male, who drove him off and seized territorial control. The new male was an indifferent stepfather, initially pecking at his stepchicks and then simply ignoring them. Meanwhile, the displaced father licked his wounds on the west side of the lake and waited for the situation to improve. Things did get better; after a few weeks of limited interaction with the female, who was preoccupied with rearing her two voracious youngsters, the new male drifted away, permitting the displaced male, who quickly regained his health, to resume parenting and rejoin his mate.

While I am delighted to see the old, evicted male pick himself up, dust himself off, and recover his position, the behavior of the usurping male on Blue-Southeast presents a puzzle. Why would an outside male hurl himself into a violent, dangerous battle, win a new and productive territory, and then relinquish it shortly afterwards without a fight? We do not understand such behavior well. However, it is possible that a young male that is physically superior to a another male with a territory can battle and defeat that male without great danger, settle and live within the territory for a time, and then decide that the breeding space does not suit his needs. I think that the preferences of loons for natal-like lakes and, later, lakes of certain clarity, might hold the key to understanding this curious behavior.

We will keep collecting data in an effort to understand why loons fight fiercely for territories and then sometimes give them back to owners. Meanwhile enjoy this video of the Blue-Southeast female enduring relentless begging by one of her chicks as she forages for it.

Two days ago we learned that the male on Hilts Lake and the female on the East Central territory on Nokomis Lake have become the latest angling casualties in our study area. The Hilts male swallowed a lure or bait and is dragging line from his bill; the Nokomis-East Central female has line wrapped around her left leg, which she now carried behind her. We are dismayed for the two loons involved, of course. The male on Hilts is among our tamest; since 2007, he and his mate have permitted us to approach them closely without alarm so that we can record an hour of their lives every week or so during the breeding season. Each of us looks forward to visits to Hilts because of the relaxed pair we encounter there. The Nokomis-East Central female too permits us to view her and her mate and young from close by. She contrasts markedly with her mate, who forces us to view from a distance of 50 meters or so to avoid getting an earful of alarm calls. When we spy a lone adult and chick from a distance on Nokomis-East Central, we keep fingers crossed that it is the docile, approachable female, not the male, who happens to be guarding and feeding the chick. (Good news: Seth reports that the Nokomis-EC female leg appeared to be dragging no fishing line yesterday, so perhaps she is out of danger.)

The list of loons ensnared in fishing tackle is longer each year. Observations and reports of this kind have become an unrelenting and disheartening drumbeat. In the early 2000s, close encounters between loons and fishing tackle were anomalies that I wondered about almost dispassionately. Each angling casualty brought sadness and frustration, of course, since one becomes attached to the loons. But such events were so uncommon that I shrugged the losses off as the inevitable consequence of habitat overlap between loons and humans. The steady increase in entanglements in recent years has sensitized me. These days I brace myself for the several – perhaps I should change that to “many” – that will occur during the season and wonder how we can assemble a team to rescue victims without compromising the behavioral and ecological goals of the research. In fact, I have begun to prod Project LoonWatch in hopes of encouraging them to put together a loon rescue squad – a group of 3-5 folks that can remain “on-call” during summers to assist birds injured by human activity.

And I have begun to worry about the loon population. Loons are clearly getting caught on fishing lines more often than before. This makes sense, as fishing activity continues to increase in northern Wisconsin. (It might also be the case that the proportion of inexperienced anglers – those likely to lose lures and fishing line in the water and not make efforts to retrieve and discard tackle that they lose – has increased.) Loons did not evolve in an environment with monofilament line, live bait and lures that mimic small aquatic creatures. Hence, they have developed no system for avoiding these hazards. If the number of loons swallowing and blundering into fishing tackle continues to increase, will so many adults die that the population cannot sustain itself?

A quirk of fishing entanglements makes the situation a bit more severe than it might first appear. As the cases of the Hilts male and Nokomis female illustrate, males and female loons are differentially affected by fishing. For reasons that we do not understand, males appear much more likely than females to ingest lures and baits, as the Hilts male did. We do not understand this difference, although one might speculate that males, having larger bodies to sustain, are less picky in chasing and consuming underwater creatures than are females. In any event, both sexes seem to be equally prone to getting monofilament wrapped around their wings or legs, but males are more threatened by lead-based tackle (sinkers and jigs) — which poisons them and kills them quickly – and by hooks that damage their throats and prevent them from opening their bills and feeding themselves. In short, males are more apt than females to fall victim to fishing tackle and in a way that is likely to kill them.

Why should we care if adult male loons are more vulnerable to anglers? Because, weirdly, males choose the nest site where eggs are to be laid and cleverly reuse nestsites where they have hatched eggs successfully in years past. So males are a reservoir of information about how breeding pairs can best use the nesting habitat within their territories to produce young. When an established, experienced male from a productive territory is replaced by new male (because of death or eviction), the new male does not know where to place the nest and must learn by trial and error. In constrast, when a female is replaced, efficient reproduction can continue uninterrupted, because the identity of the female has no impact on nest placement.

Thus, I grieve for the Hilts male not only because he is a sweet bird who permits us to watch him without complaint but because his demise will cause a new, ignorant male to settle, who must blunder from one failed nestsite to another before he discovers a good one that he will use year after year. And his years of inept breeding will subtract many offspring from the population.