Recently, I made the kind of finding that gives scientists fits. It came about in the same manner that findings often do initially: a hunch.

Since I spend much of my life either working closely with loons or poring over data that describes their breeding success, I am in a good position to notice subtle changes that occur over time. Occasionally when out on a lake, I observe a breeding event and think, “Wow…..that did not happen in the old days!” Then I retreat to my computer, look at data from years past, and see if I am correct. I have to confess: in many cases, I am wrong.


This past August, I noticed what I thought might be a growing pattern in breeding ecology. Mated pairs, it seemed to me, were less often rearing two chicks to fledging. That is, they were either hatching one chick and rearing the singleton only or hatching two chicks and losing one. At least that was my impression. In this case, field data confirmed my suspicion, as the above graph shows. The proportion of singleton broods has risen during the study. In three of the past five years, in fact, two-chick broods were quite uncommon, making up only 1/5 of all broods. Most of the pattern, moreover, appears to result from failure of one of two eggs to hatch, rather than loss of the second chick after hatching.

Faced with a puzzling and unexpected finding, I looked immediately at the usual suspects. Black flies, which have also been worse in recent years, are an obvious possibility. Flies harass incubating loons, reduce incubation times, and might reduce hatching success. In fact, I was convinced when I wrote our recent paper that black flies were the culprits. But then 2018 happened. This season featured a warm spring, a rapid die-off of flies, and very few fly-induced abandonments. So we could not blame flies for the low hatching rate of eggs in 2018.

What about the changing climate? As I have emphasized in a recent post, warm weather is projected to drive loons northwards; could it also be the root cause of the lower hatching rate? I looked to see if warmer May temperatures are correlated with reduced hatching, but they are not. (In fact, warmer temperatures are associated with a slightly higher hatching rate.) Likewise, precipitation might, in theory, reduce hatching rate. Again, years of higher May rainfall were not years of lower hatching success. I breathed a sigh of relief to learn that the lower rate of hatching does not (preliminarily) appear to represent the harmful leading edge of climate’s impact on loons.

Two possibilities remain. First of all, there is a small chance that the pattern is a statistical anomaly — that hatching rate is not actually falling, but that the result occurred by sheer chance. Scientists must always be circumspect about their results, and the statistical test says that there is a 0.6% chance that the finding does not represent a true pattern. (That is roughly the likelihood that you toss a fair coin 9 times in a row and get “heads” every time.) Second and more likely, some unknown factor is at work here. Might there be an environmental contaminant, picked up by loons, that increases developmental abnormalities in embryos or perhaps causes adults to cease incubation of the second egg prematurely? Might disturbance of incubation by humans be the cause of lower hatching success? These possibilities — and numerous others — generate testable predictions, and I will test them. In the meantime, let’s all keep our fingers crossed that the distinct decline in loon hatching success over the past 20 years is, after all, just a blip.




Humans are not good at thinking about the distant future. We are not alone in our short-sightedness. Living things, in general, are obsessed with the here and now and oblivious to what lies far down the road. There is a very good evolutionary reason for focusing on the present. Animals that succeed at surviving and protecting their progeny leave more young than other animals (in this case, hypothetical ones) preoccupied with what conditions might be like for their grandchildren and great-grandchildren. Animals that attend to their own survival and that of their offspring simply leave more offspring. Thus, natural selection can be said to favor animals that focus on the present — and animals within natural populations are chiefly descendants of parents and grandparents that cared for their own survival and that of their offspring. The short-term view makes sense evolutionarily.

Our very logical tendency to heed the here and now at the expense of the future has a limitation. Focus on the present adapts animals well to a stable environment, but leaves them poorly adapted to an environment that changes rapidly. Over evolutionary time, environmental change has generally occurred slowly enough to cause little problem for animals that live only for the present.

But humans have hastened environmental change. Anthropogenic changes have taken many forms, including introduction of invasive species, environmental degradation, and wholesale alteration of landscapes and vegetation. Perhaps surprisingly, many non-human animals have been able to keep pace with human impacts. In fact, some — crows, gulls and raccoons come to mind — have benefitted enormously from human activity. Others, of course, have become extinct, endangered or have seen their geographic ranges contract because of humans. We could quantify human impacts of each and every non-human species, if we cared to, and place each on a chart from least- to most-impacted.

Where on the chart would the common loon fall? Considering that loons are often viewed as the “voice of the wilderness”, one might suppose that they would suffer greatly from human encroachment. In fact, loons are hanging in there better than many other vertebrate animals. Knocked back in the middle of the 20th century, the common loon population has rebounded. Breeding populations are now generally stable or even increasing across most of the northern tier of United States. My study area in northern Wisconsin is typical; loons have re-colonized many lakes in the past few decades from which they had retreated. So loon populations are thriving despite extensive shoreline development, entanglements with hooks and fishing line, and increases in methylmercury levels, among many other challenges.

A new anthropogenic threat now looms that is more extensive and unrelenting than others that loons have faced. Climate change has already caused many geographic ranges of North American animals to recede northwards. A recent study showed that bird species differed greatly in their northward shifts, but that, on average, breeding ranges are marching northwards by over 2 km per year. We have a bit of an apples and oranges problem here; the bird species included in the study varied greatly in their diets and habitats. Some, no doubt, are highly dependent upon temperatures (and related factors like vegetation) for their survival; others are not. So it is difficult to project precisely how the geographic range of the common loon might be affected. But do this: take a look at Audubon’s animated depiction showing the contraction of the loon’s breeding range.

Two patterns are immediately clear from the animation. First, the northern Wisconsin loon population (and abutting populations in Minnesota and Michigan’s Upper Peninsula) exist on an isolated “finger” that projects southwards from the heart of the range, which lies in Canada. Second, the model paints a very bleak picture of the future loon population in northern Wisconsin. According to the model, loons are projected to be much less abundant in northern Wisconsin by 2050 and gone altogether by 2080.

Now, a word of caution. Audubon scientists have attempted to distill the climate down to two main factors: temperature and precipitation. On the basis of these two climatic factors, the current distribution of the species relative to these factors, and the projected future climate based on the report of the Intergovernmental Panel on Climate Change (IPCC), they have produced theĀ  animated graph that loon enthusiasts like us find so disturbing. Their projection is likely to provide a crude estimate of the impact of climate change on loons, not a precise one. That is, loons are likely to cope with climate change better than most other birds — as they have other environmental threats. Then again, loons might be especially sensitive to climate change and retreat northward more rapidly than the study predicts.

Like many other humans, I am obsessed with the day to day. I have studied loons as if they would be around forever. I have battled to obtain grants to keep my study afloat, to publish my papers in high-impact journals, to hire diligent field technicians who would collect reliable data. Now, forced by changing environmental conditions to glance towards the future, I can scarcely believe that the animals that I have learned so much about and grown so fond of might be well on their way to vanishing from Wisconsin in my lifetime.




In my last post, I told only half of the story of the explosion of the Cunard family — the cheerful half. If you read that post, you know that, following the eviction of the Cunard male and abuse or neglect of the chicks by the evicter, one chick made a daring 1/4-mile trek across land to Hasbrook Lake and is now happily ensconced in that loon family.

The other Cunard chick was not so fortunate. In fact, following interviews of campers and the camp steward, we now know that a day or so before the eviction that led to the desperate dash of one chick to Hasbrook, its sibling had swallowed the live bait and hook used by a camper. As he described it to me, the fisherman panicked and did what most do when they have hooked a loon: he cut the line. I discovered the aftermath of this hooking. The thoroughly consumed remains pictured above suggest that the injured chick became weak, took refuge on shore (as seriously injured loons do), and was attacked and killed by an opportunistic mammal or scavenged after death. Its four leg bands confirmed its identity; the threaded line and fishing snap I found confirmed the cause of death. So the eviction that occurred on July 30th and 31st was actually the second unfortunate turn for the Cunard pair during the last three days of July.

We have discovered several such hookings during our study, despite the fact that anglers do not trumpet them. Perhaps we should take a moment to describe what to do when a loon takes your hook. The best outcome is removal of the hook by the fisherman. Removal of the hook gives the loon a good chance to survive the encounter. Cutting the line, on the other hand, frees the angler but leaves the hooked loon with a death sentence. A hooked loon (or other animal) on the end of a cut fishing line has to contend with a hook or lure that it probably cannot dislodge on its own. Its feeding impaired or prevented altogether, a hooked bird will probably succumb to starvation or predation resulting from its weakened condition. The second best outcome is to cut the line and immediately inform a local wildlife official of what has happened so that he or she can get help for the bird. In many cases, a hooked bird can be captured and de-hooked by me or someone else trained to do so. In other words, if you cut, don’t cut and run. Those of us who study and love loons will do our best to save one that is in trouble.

The Cunard chick’s death is a case in point. Had we known about the hooking, we would have had little difficulty re-capturing the chick and likely removing the hook as well. In that case, Hasbrook Lake might have ended up with four chicks rather than settling for three!

To those of us accustomed to looking at loons during the summer, the sight of an adult caring for three chicks — as in Laura Unfried’s photos from two days ago — is peculiar. Loons, of course, almost always lay only two eggs. If they are lucky, two chicks hatch. It is by no means certain that those two chicks will survive to fledging age. In fact, 47 of 61 breeding pairs we study have one chick, not two. So the spectacle of two adults caring assiduously for three young was startling.

Close inspection of Laura’s photo from Hasbrook Lake reveals another peculiarity: the central chick is much larger — perhaps 10 days older — than the other two chicks. (Note that the left-hand chick is entirely downy with a small bill, whereas the center one has the anterior part of its head coming into adult feather and has a bill nearly as thick as the parent’s.) The obvious size disparity told Eileen Lonsdorf, who reported the third chick three days ago, that her nicely balanced family of two parents and two chicks had been joined by an interloper.

How could a huge, healthy chick somehow get separated from its biological parents and join another family? Territorial behavior among loons guarantees that each breeding pair will nest and rear its young far from other pairs. The likelihood of a chick straying from its own family to join another — fortuitously or by design —


seems remote.

One lesson that you learn if you do something for a long time is that rare events do occur. Chicks do very occasionally leave their parents and territory to join other families. We have noted two causes for such chick dispersal. First, starving chicks, especially beta chicks on small food-limited lakes that are being physically beaten by their alpha siblings, sometimes attempt to escape the abuse and find an alternate loon family nearby that will feed and protect them. Second, chicks that lose one or both parents to territorial eviction are forced to flee their natal territory and seek parental care elsewhere, if the adult that evicted a parent physically attacks them.

Solitary journeys by displaced chicks seeking new homes are desperate enterprises. One reason for this is that many lakes with loon chicks simply have no neighboring pairs with their own chicks that might be joined. Even if a displaced chick is fortunate enough to find a nearby pair with chicks, they are likely to be much older or younger than itself. If so, it is unlikely to be accepted by the new family. I vividly recall a case in 1999, when an abused beta chick undertook an astounding 1/2-mile trip across woods and roads from Benedict to Bug Lake in Vilas County, only to land with a foster sibling three times its size that beat it mercilessly until it perished.

Since the monster chick that joined the Hasbrook pair is a robust, well-fed individual, we could rule out that it fled to Hasbrook because of sibling abuse or lack of food. So we were left to conclude tentatively that a nearby territorial eviction forced this young loon to relocate. We pulled out a map to assess the possibilities.

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Immediately, we pinpointed Cunard Lake, which is separated from Hasbrook by a quarter mile of woods and bog, as the likely source of the wandering chick. Cunard, a regular study lake of ours, had two large, healthy, 5-week-old chicks at our last visit on July 25th. Yesterday, however, I found the lake empty of loons, except a single floater adult. The steward of the campground reported that loons had been chasing each other repeatedly across the water on July 30th and 31st, which indicated a protracted territorial battle. The absence of the territorial pair suggested strongly that the breeding male had lost the battle to a usurper and either died or been forced to abandon his territory and chicks. This tragic event, in turn, would have scattered the rest of the family and subjected the chicks to attacks by the new male owner.

Last night we captured the peculiar but close-knit family of two adults and three chicks on Hasbrook. It will require genetic analysis to be certain that the huge new chick on Hasbrook is a refugee from Cunard and the offspring of the displaced Cunard pair, as we surmise. But we have strong reason to believe that he is a most fortunate survivor of a desperate overland journey.

Few loons have endured the frustrations that the current Mildred female (“Taupe Stripe”) has. Initially captured and banded on Soo Lake in 2004, she reared two healthy chicks with her mate in that year. But each year since has yielded no offspring for this bird, despite consistent effort.

Taupe Stripe’s struggles started when she was evicted from Soo by a stronger female in 2006. Thus began an itinerant lifestyle: temporary settlement as a loner on Goodyear Lake and frequent intrusion into other lakes in the area, probing for an opening. In 2010, she finally secured and defended a breeding position on Maud Lake. Like other small, shallow lakes, though, Maud suffers from a limited prey base. Efforts by the loon pair there in the three years before her arrival were excruciating — hatched eggs followed by starvation of chicks in the 2nd week of life, the first week, and then the fourth week. Taupe Stripe and her two mates fared no better, failing to produce young in 2010, 2011, 2012, and 2013. She fell off our radar in late 2013 and 2014, but resurfaced in 2015 and 2016 — again, as a floater.

When day to day survival is a challenge, as it is for loons, you focus on the present. Taupe Stripe ultimately settled on Mildred Lake in early 2017. Unlike Maud, Mildred is a large, clear lake with a strong record of fledging chicks that hatch. Although she and her unbanded mate lost a small chick on Mildred last year, they have turned things around. Last week we caught Taupe Stripe, her mate (now banded), and their strapping six-week-old chick. Things were still looking good for the family upon my visit there today. And thirteen years of pointless wandering, frustration, and disappointment are forgotten.

For the past several years, I have begun to turn my attention to the effect of lake size on breeding success of common loons. Clearly loons on large lakes produce more and healthier chicks than those on small lakes. We showed that on our paper from six years ago. This raises the question of why loons ever attempt to breed on small lakes. They are doomed to failure — or at least to greatly reduced likelihood of success. The water is muddied further by the fact that loons reared on small lakes prefer to breed on small lakes themselves. That’s right: chicks fortunate enough to avoid starvation on small, food-limited lakes replay the whole scenario as adults, subjecting their young to the same travail they themselves faced.

The puzzle of loons breeding on small lakes was thrown into start relief again yesterday, when I visited the Wind Pudding-West territory on a scouting trip for nocturnal capture. After not finding the chick and parents in the shallow bay on the lake’s western side, where we had seen them on previous visits, I headed towards the channel that connects that bay with the main lake. I was crestfallen when I reached the channel, as it was choked with lily pads and grasses to the point where it was difficult for me to find a passage through — even in a canoe. This discovery led me to doubt whether the pair might lead their chick through the channel and into the main lake as a means of finding more food for it. I began to fear that — walled off from an abundant source of food — the chick had probably starved to death on the shallow western bay since our last visit.

I had underestimated the determination of the pair to provide for their chick. As I paddled to the end of the navigable portion of the channel, I heard a chick’s desperate cries to its parents. I spotted the chick about half way across the marshy isthmus that now separates the shallow western bay from the main lake. The isthmus is no more than 20 meters wide perhaps, but it is densely overgrown with marsh grass to the point where the chick — equipped only with legs at the very posterior of its body — was forced to lunge awkwardly forward in order to make headway towards the main lake. To make matters worse, the chick had no clear idea of where it was going. I caught the chick’s initial confusion on video, as it sits within the marsh grass, uncertain how to extricate itself.

The second installment shows the chick after it has blundered around in the grass for a time but finally gotten a sense of where its parent wants it to go. The chick stops to give a distress call, then hears its parent call to it, which seems to give the chick the strength to complete its journey. (Apologies for the nervous narration and grainy video!)

The fact that the parents can entice this year’s chick to cross the isthmus and take advantage of food in both the western bay and main lake means that they are better off than they were here in 2016, when a chick wasted away and finally died of starvation. However, we caught the chick and female in the video last night. They are both severely underweight. The female, in fact, has the lowest mass we have ever measured for an adult loon. So even if the pair can find enough food to fledge the chick, chick-rearing seems to have taken a toll on the parents. Add yet one more item to the growing list of reasons to avoid breeding on small lakes with limited food.

The deformity was obvious back on May 17th, when we first saw her. The breeding female on Johnson Lake had part of her bill jutting upwards at a crazy angle. At first, viewing her at a distance, we thought that the dark spike that appeared to emerge from her bill might be a lure of some kind that she had latched onto mistakenly and been unable to dislodge. But, as Elaina’s photo shows, the distal half of her bill is bent upwards at a 45 degree angle. We were alarmed at her situation, which appeared uncomfortable, at least, and deadly, at worst.

But she behaved normally. Since we carefully observe, painstakingly describe, and publish articles about loon behavior, we habitually assess any loon’s comportment that we see according to thousands of others seen before. Her diving and foraging was normal. Far from permitting too-close approach by humans — a common red flag that can indicate severe injury — she was actually rather skittish. She pointedly moved away from us whenever we approached in an effort to look for leg bands.

Anatomically, she is far from normal. Bird’s bills consist of a matrix of bony support, covered by a keratinized epidermal layer (rhamphotheca). In other words, the displaced part of this loon’s bill should comprise not merely soft tissue but bone. Mark Pokras, associate professor emeritus at Tufts School of Veterinary Medicine, assures me that the fact that the bony foundation that should extend to the bill tip is missing means that it “will never grow back normally”. The best we can hope for, he says, is that the bent keratinized tissue — all that remains of the end of her upper mandible — drops off eventually. I was chagrined to hear this news but heartened to learn also that Dr. Pokras has, during his decades of loon anatomical study, seen about 10 cases where large portions of loon bills have been missing. These cases include a male in Maine that had only half of an upper mandible (as this female does) but that fed itself normally, held its territory, and produced offspring in multiple years.

That loons can survive an injury of this kind to a crucial feeding organ and still breed seems remarkable. I suppose their resilience might be explained partly by theĀ  challenges they face routinely across the range of different landscapes they inhabit. That is, an animal that must locate, pursue, and capture a broad spectrum of actively-swimming prey — in water that is sometimes fresh, sometimes salty; sometimes clear, sometimes turbid, must be a flexible and adaptable creature indeed.

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.