I do not know why I am still surprised by it. Again and again we see territorial females vanish, only to reappear as intruders in their former territory or another in the same neighborhood. The theme has been consistent throughout my quarter-century-long research project on loons. In fact, it is about a 50-50 proposition that a female who disappears from her territory will be resighted nearby in the next year or two.

I suppose the reason why I am always wrong-footed by female reappearances is that I am calibrated to human social conditions. Unlike loons, humans are highly social and benefit from a vast network of family ties and friendships. Humans have evolved to cultivate those bonds — leaning upon family and friends when necessary; providing support, in turn, when called upon to do so. Humans, therefore, rarely vanish without a trace from an area where they have lived for many years. We typically maintain most or all of our social connections even when we shift from one location to another.

Loons, in contrast, are quite alone. To be sure, an adult male or female has the companionship of its mate for several months each year. But this companionship is ephemeral and conditional. When a loon’s mate is evicted by an intruder, the loon faces a stark choice: remain with its mate of many years but move to a new territory, or stay on its territory and accept the evicting intruder as its new mate. Adult loons always spurn their long-term partner and make the coolly pragmatic choice.

Knowing loons as I now do, I should not have been surprised when Linda sent me this crisp photo of an intruder departing from Muskellunge Lake. Although the bird is missing a colored leg band from its right leg, it was not difficult to determine that this intruder is the former female from Manson Lake, who has not been seen since fall 2015. Since loons almost never desert a territory, we can surmise that “Silver over Red, Mint Burgundy over White” (as we affectionately call her) was evicted by the 9-year-old female from Rock Lake in Vilas County who was first seen on Manson in April 2016 and still owns that territory. The eviction forced “Silver over Red” into an itinerant existence. Though you might think that a veteran of 12 years of breeding and mother of 11 fledged chicks had earned better treatment, this 20+ year old now spends her time drifting from one inhabited lake to another as an intruder, searching for a new mate and territory so that she can resume breeding.

It is tempting to pity Silver over Red. These past two years have been difficult and dangerous for her, no doubt. But rather than viewing her and others like her as individuals whose best years are behind them, I always feel an intense admiration for displaced female loons. While many of us humans respond to setbacks with a shrug of acceptance, female loons never stop hunting for a new opportunity.

Science is a cumulative pursuit. That is, the thirst for knowledge is never fully quenched. Rather, we answer one question, only to reveal another puzzle or two more. And thus begins another search for answers. That is certainly how my 24-year-old loon project has gone. But sometimes we reach a point where a vexing question is finally laid to rest, and it feels as though we have made real progress. I am at such a point now.

Let me back up. If you have been following this blog, you know that senescence in old loons is a phenomenon we have recently discovered. Senescence, loss of body condition and decline in survival rate in aging individuals within a species, is all too familiar to me and other humans. During the past twenty years, many studies have reported senescence in birds, mammals, fish, and reptiles. So what? Well, we expect that animals that lose condition as they grow old will change their behavior in response. In other words, scientists have long predicted that senescing individuals should start to behave so as to leave more to their offspring and care less for themselves. To put it another way, old individuals should be willing to take a hit to their survival if it allows them to pour more resources into their young and help their young survive. This makes sense, of course, because old individuals reach a point where they stand little chance of surviving longer, so they would do well to give whatever they can to their offspring, which DO have a bright future. Animals that behave this way should leave more and healthier offspring, and thus this behavior should spread in populations. This very logical idea is termed “terminal investment”. Again we can all probably think of human parallels.

Terminal investment, which I have mentioned before, has become a central theme of the loon project, ever since we published a paper 9 years ago on fatal fighting of males. Terminal investment became interesting to us because it was the most plausible explanation for such lethal contests. Our reasoning was as follows. If males are willing to die to defend their territories, then they must reach an age at which they have little to lose. And if males have little to lose, this must mean that senescence hits males (but not females, which seldom battle to the death) very hard to the point where old males have little future to look forward to. In this case, it might make sense for them to fight like crazy to hold a territory for another year or two, rather than give it up easily and leave themselves nowhere to breed during their last year or two of life. So we have two clear predictions here: 1) males, but not females, must start to die off at a certain age, and 2) males beyond this age must still fight like hell for their territories. It is this clash of body condition and behavior among old males that might cause fatal fighting.

At the time we started to consider the terminal investment hypothesis as a means to explain reckless battling by males, we had almost no solid information on the ages of males in our population. With patience and tireless field work by dozens of us, we have now turned things around. Analysis of loons of varying age has shown us that many males hit the wall at age 15. First, and most important, they start to die at a high rate. You can see from the figure below that males (blue bars) are suffering higher mortality than females (red bars), whether they are on territory (Terr) or without one (floaters: “Float”).

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But males also lose mass at age 15, indicating loss in body condition, as shown here:

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Finally (and predictably), males get evicted from their territories at a high rate at age 15:

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Wow, males are really getting slammed after they pass the age of 15 years!

So all of these data tell us that the first prediction of the terminal investment hypothesis, abrupt senescence at a certain age in males but not females, is clearly met in loons. That age, surprisingly, is only fifteen. Females clearly remain strong, healthy, and vigorous well past age fifteen.

As hard as the first prediction of terminal investment was to test, the second prediction is even harder. You see, fights are common in loons if you take the perspective of a loon’s lifetime, but they are quite uncommon if viewed from the standpoint of human observers in canoes. In other words, most individual loons have engaged in several major battles during their long lives, but territorial battles are not common during day to day observations and often occur so quickly that we are not present to witness them.

Patience pays, however. Since we can draw upon 24 years’ worth of field observations, we now have a trove of observations that we can search for any evidence of aggression and territorial behavior. I made this search, looking for two kinds of evidence: 1) territorial yodels, which serve to communicate a male’s aggressive tendencies and willingness to battle, and 2) out and out aggression, in the form of battling, lunging, chasing and underwater attacks launched by territorial loons on intruders to their territories. I was simply asking “Do old male loons (above age 15) tend to maintain a high level of yodeling and aggressiveness towards intruders?”.  The answer is a resounding “Yes”:

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As you can see from the figure above, old males actually increase their tendency to yodel (yodels per intruder), compared to young males. Similarly, old males step up their aggression (see below) and contrast in this way with females, who show no increase:

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By the way, all of these patterns I have shown are “statistically significant” via tests that I have performed.

You cannot be as excited as I am about this set of results. No one is. But, as I mentioned, this is one of those rare cases where we have finally managed to answer a burning question to our satisfaction. Even better, the question is one that had been the foundation of my research funding from the National Science Foundation. So I can now report to them that I have found the holy grail! What makes this clear finding even more significant is that terminal investment is quite rare in vertebrates. Of the hundreds of species studied thus far, the only other one to show such a clear pattern of terminal investment is the California Gull. Appropriate, don’t you think?

(Photo by Woody Hagge.)

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It’s like deja vu all over again at Muskellunge Lake this week. Thanks to Linda Grenzer’s vigilance, we have learned that the productive and photogenic pair on Muskellunge came back together this week — just as they did about this time in 2014! As Linda’s photo shows, “Clune” and “Honey” (Linda’s names for the banded pair) were a bit wary of each other at first. This shyness should not be surprising, as they have both lost their breeding plumage and donned winter attire. According to Linda, though, they rather quickly recognized each other and paddled off contentedly together. Apparently after 6 years of being together, and rearing of 10 chicks to fledging, familiarity with your partner involves more than just feathers!

We have very poor information on the behavior of territorial loons in the fall, so we can only speculate about what it means that an established pair seems to come together routinely just before fall migration. As a behavioral ecologist, I am inclined to interpret such meetings as more calculating than romantic. I cannot resist the temptation to view such a meeting as a final patrol of this precious breeding lake by its owners to ensure that no intruders have tried to put down roots in hopes of claiming Muskellunge in 2017. To any such pretenders, the presence of a tight pair late in the fall would signal that the territory was not available without a costly battle. But that is just speculation.

I must apologize for taking so long for this post; I have been busy exploring loon survival patterns. Indeed, I have been able to sharpen our view of senescence considerably. Some of you will remember that I announced several months ago that males senesce, while females do not. While I was excited about the apparent finding, it seemed to defy logic and begged for elaboration. With the return rate data from 2016 in hand, I was able to reanalyze survival patterns of both males and females banded as adults on territory. As is becoming all too common, I must eat crow. It is still clear that males senesce — those between 5

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and 19 years of age survive at a rate of 94% annually, while those 20 years and older survive at an estimated rate of only 71% (see graph, above). That is a steep drop-off in survival rate, indicating clear senescence. But the newer, more robust dataset has shown that females are not immune to the march of time. Indeed, 5 to 19 year old females survive at about 94%, like males; 20+ year old females fall off to 79% survival (see above graph). It is a tad deflating to realize that the world was more complicated than I had reported before, but also a relief to know that — and don’t take this wrong — females are not immortal!

One more tidbit relating to senescence. I have been able recently to complete a survival analysis of loons that were banded as chicks in the study area or nearby. (We call these birds “ABJs”, or “adults banded as juveniles”.) The most striking outcome of the ABJ survival analysis is, again, a difference between females and males. In this case, male survival plummets from 94% at ages 4 to 14 to 78% at ages 15 to 18 among both territory holders and floaters (adults without a

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territory; see graph above). In contrast, females show steady survival of about 92%, regardless of age or territorial status (see above graph). Males, in addition, suddenly begin to lose their territories to eviction at the rate of 38% when they turn 15 years old, while females only suffer eviction at a rate of 12% from 4 to 18 years of age. So here we have evidence that males not only die at a higher rate once they turn 15, but that they also become vulnerable to being booted off of their territory.

If you have paid attention closely through the litany of data I have dumped on you, you have probably noticed one final pattern: the ABJ analysis showed senescence occurring in a younger age class than did the analysis of territory holders! That is, male ABJs senesce at age 15, while male and female territory holders show no senescence until age 20. This is a brand new finding that I am still puzzling over, but there is an obvious explanation. Territory holders are likely to comprise a fitter class of adult loons that have shown their ability to fight for and defend territories and also rear chicks. In contrast, ABJs are only chicks that have survived to adulthood and returned to the study area. They are not “battle-tested” by having claimed a territory and bred there. Much later senescence among the evolutionarily fittest set of loons in the study area should not surprise us.

I am not ready to stake my reputation on this latest finding, but if it holds up, it dovetails nicely with one of the long-standing debates among ecologists concerning territorial breeders and floaters. We have long wondered whether territory holders should be regarded as floaters who got lucky and found a territory, or whether the two groups are, in fact, distinct classes in terms of fitness. The steep drop off in survival of male floaters in loons, compared to male territory holders, would strongly support the latter interpretation.

If you have been floundering lately, as I have, let’s gain some perspective by considering the plight of the Silver Lake loons. The pair’s struggles began in mid-May, as black flies thwarted their efforts at incubating a first clutch of eggs. According to Pat Schmidt, who watches the pair carefully throughout the breeding season, incubation proceeded normally during the nighttime — cool temperatures kept the relentless pests at bay. But the marked female and male were unable to stay on the eggs during daylight hours, when black flies were active and biting. On again, off again incubation finally gave way to abandonment during the last week of May, but the pair reset themselves quickly, adding two additional eggs to the two they had earlier tried and failed to hatch. Despite the cumbersome task of warming four large eggs simultaneously, the birds produced a chick at the very end of June. Their fortunes seemed to have turned.

The greatest risk faced by a breeding loon pair with a chick is our national bird. Bald eagles nest on tall white pines along lake shores and are a frequent sight over lakes. Indeed, eagles are such a routine part of the scenery on the lakes that loons often deign to wail at them as they pass overhead. Eagle fanciers might try to convince us that these raptors even purposely lull loons into a false sense of security with their constant, mostly innocuous flights nearby so that they can occasionally strike at loons suddenly with deadly purpose. An opportunity for such a surprise attack might occur when an eagle appears just above the tall trees at the lake’s edge as a week-old chick’s parents both happen to be underwater diving for food. Perhaps it was such happenstance that allowed an eagle to carry off the Silver Lake chick on July 2nd. In any event, eagle predation brought the breeding efforts of the pair to an unsuccessful close this year.

The sting felt by lake residents at the loss of the chick had begun to abate by July 18, at which point the territorial female, “Copper” (named for one of her plastic leg bands), found herself in a desperate battle. She was beaten badly, chased across the water, attacked from below as she rested on the lake surface, and finally forced to take refuge on land to escape further damage. By the time the violence had ceased, Copper had to be carried, helpless, to the Northwoods Wildlife Center. She died there a few days later.

As I have made clear in numerous posts, males are the ones that battle dangerously (apparently because of senescence) in most cases. So how do we explain the latest Silver debacle? An oddity concerning contestants might offer a clue in this case. Copper, who had reared chicks on Silver in 2014 and 2015, had battled repeatedly for ownership over the past several years with her bitter rival, “Mint”, the previous Silver female and mother of the chicks in 2010 and 2012. Even after losing the territory to Copper in early 2013, Mint was a frequent intruder into Silver Lake. Hence, both females had raised chicks in multiple years with the male, and both were highly motivated to vie for control of the territory. In addition, banding records indicate that Copper and Mint were of very similar size.

Now to game theory. If an animal encounters a long-lived opponent with which it is very closely matched in fighting ability but happens to get the upper hand at some point, it might then pay for that first animal to press its advantage and even kill the opponent. Why? Because our research has shown us that closely-matched pairs of females, like Copper and Mint, often give each other fits. Two females on Heiress and two others on Oscar-Jenny were so close in fighting ability that they traded off ownership of those territories over many years, each female hindered in her breeding efforts because of the constant interruptions of the other. The result was poor reproductive success for both rivals. Although there is enormous risk involved, it might occasionally pay for females to exhibit the vicious battling we associate with males. Specifically, a lightning strike to finish off your archrival might sometimes be worthwhile to avoid a chronic, destructive feud.

 

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In many recent posts, I have emphasized a certain theme: male loons begin to die off at a rapid rate after age twenty, while females linger on. Part of the reason for this contrast is the nature of territorial contests in each sex. Territorial males fight hard in attempting to hold their breeding position on a lake and commonly die in territorial battles. With rare exceptions, territorial females survive eviction from a territory, move to an unoccupied lake nearby, and resettle on a new territory when opportunity permits.

While the escalation of male territorial battles is interesting in itself, it also impacts the composition of the breeding population. Specifically, adult male loons’ propensity to die frequently in battle skews the sex ratio towards females in the breeding population.

These excess females are “floaters” — adults capable of breeding but prevented from so doing by the lack of a mate and/or a territory. Floaters are the loons that one sees living alone on small lakes, drifting about aimlessly on large lakes, and intruding into territories from time to time to confront breeders. A large proportion of the loons that gather in flotillas of five adults or more during July and August are floaters. Floaters can be thought of as “hopeful breeders”; that is, they are always ready to settle and breed with a mate and territory, if they can find one. The excess of female floaters means that there are always far more of them looking to settle and breed than there are male floaters able to pair with them. In effect, males are snapped up by females as soon as they become available for breeding.

In May of this year, we re-encountered one of our veteran breeders, “Silver over Blue, Green over Orange” (or “S/B,G/O”), whose breeding history illustrates the striking contrast between males and females brought about by male-biased mortality. S/B,G/O was first captured and marked as an adult in 1997 on Dorothy Lake, where she raised two chicks with her mate. Her mate was evicted in 2001 and died either during eviction or shortly afterwards. But she lingered on. When an opening became available in 2002, she settled and nested with a different male on Hasbrook Lake, just a few miles to the northwest. Having failed to raise chicks on Hasbrook, S/B,G/O (now at least 14 years old) evicted the female breeder on Hodstradt in 2004, paired with a third male, a six year-old, and reared four chicks there during the next three years. She followed this young male to Horsehead Lake in 2008, when he was driven off of Hodstradt, and the pair fledged 3 more chicks over the next four years on their new lake. When the male was evicted yet again in 2013, S/B,G/O traded experience for youth and found a new six year-old male as a breeding partner. We breathed a sigh of relief when she broke up with this youngster after a year together, as he was unfortunately her son from Hodstradt! Then 23+ years old, S/B,G/O again became a floater, forced to return to the breeding grounds in 2014 and 2015 with no clear prospects for breeding.

I have become attached to the birds in the study area, so I was delighted to find S/B,G/O back at Hodstradt in May of this year with her fifth recorded mate. At 26+ years of age, she is perhaps fortunate to be paired again. Her mate this time: a four year-old hatched on Clear Lake. We observed no breeding attempt by this new pair – only a small percentage of four- year-old males that settle on territories actually nest – but it is likely they will nest in 2017.

As a human, I like to think of S/B,G/O’s life as a lesson in resilience – the dogged refusal of an animal to forsake breeding despite repeated setbacks and advancing age. But, as a behavioral ecologist, I think of this female more as a striking example of how animals adapt to maximize their breeding capacity regardless of the breeding environment they face. By the way, S/B,G/O is not the only female in our study area who has continued to breed despite frequent changes of partner. S/R,O/O, another 26+ year-old from Swamp Lake that we recaptured a few nights ago (see photo with Eric), has gone through at least 5 younger mates during her 20 years of breeding there. Clearly the pairing of tough, old females with much younger males is – as my daughter says – a thing.

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He doesn’t look it, but this male from Townline Lake, just outside of Rhinelander, is at least twenty-seven years old. He is among a dwindling few males from among those we banded in the mid 90s. This bird was banded in 1994, at which point he was certainly at least five years old, which means that he was hatched in 1989 or before. Thus, twenty-seven is a minimum estimate for his age.

The age of “Silver over Red, Orange over Green” (as I call him affectionately) is not his only remarkable attribute. What sets this individual apart from most others is his ability to hold onto his territory year after year while fledging healthy chicks. (Below, he relaxes near his mate and two strapping chicks from 2016.) A successful common loon is not only good at locating safe nest sites and defending and feeding young. A breeder that wishes to

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reproduce successfully must confront intruders that land in the territory without warning throughout the breeding period.

Intrusions are especially frequent during the chick-rearing period. A common scenario plays out as follows. Early in the morning, a male is diving for food, while his two chicks track his progress from the surface. Each time he surfaces, the chicks rush over to him, snatch food from his grasp, and nibble relentlessly at his bill, neck and chest, signaling their unquenchable appetites. On one occasion, he surfaces holding a small yellow perch, only to find five adult loons in flight above his lake. He drops the fish, gives a short barking call, and the chicks dive and head to the nearest shore. The male too dives but surfaces near the middle of the lake, drawing the now-descending intruders to himself. Three quarters of an hour later he has driven off the intruders, thanks in part to a lunge and point yodel that caused his five visitors to scatter and tremolo. Shortly afterwards his mate returns, and both parents forage for the chicks. The family suffers no further disruptions until the evening, when another group of three nonbreeders circle and land, causing yet another brief skirmish.

Considering that a large pool of territorial intruders are constantly sizing up the resident male or female of any successful territory for an eviction attempt, it seems remarkable that residents are able to hold on to their territories for even a single year. Yet Silver over Red, Orange over Green has put together a string of 23 years of straight ownership, the only blemishes a half-year in 1996 and another in 2003, when he was briefly deposed. He has fledged 20 chicks during his breeding career with four different mates. This male is not the only resident with an impressive resume. A female on nearby Langley has fledged 17 chicks on that territory since 1995, while the O’Day female has been on territory since at least 1997 and has produced at least 16 full-grown chicks during her breeding career.

But female loons are survivors. Females enjoy a high rate of survival and no detectable senescence well into their twenties. Males, on the other hand, hit the wall abruptly at age 20; almost half of all territorial males of age 20 will perish before the subsequent year. So when we see a male who defies the odds, like this one, it is worth looking closely to see if he possesses an attribute that sets him apart. As a scientist, I am loathe to draw conclusions based on a sample of one. Colleagues in my field would dismiss any such conclusions out of hand. But today Nelson, one of my Chapman research students this year, reported that Silver over Red, Orange over Green is the tamest bird we have ever measured in the study area. So let me invite ridicule by advancing a very preliminary hypothesis. Perhaps the key to lifetime productivity in a habitat rife with human recreation is picking one’s battles carefully. Maybe by ignoring the inquisitive, well-meaning primates in their watercraft, this male has been able to conserve his metabolic resources for provisioning young and driving off pesky intruders.

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!

 

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.