In Linda‘s last round of photos from Bear Lake, you can see the numbers clearly. At the narrow end of the metal band, a fuzzy, curling “6” followed by a clear, swooping “2”. Those two digits — either one of them, actually — identify the Mystery Female of Bear Lake as the chick we banded on July 18, 2005 on North Nokomis Lake. This photo culminates several days during which Linda patiently stalked the female with her camera lens until the bird finally pulled her metal band out of water in an orientation that permitted Linda to photograph the two digits we needed to see from the nine-digit number code.

The story of the Mystery Female concerns more than just solid detective work and crisp photography; it relates to the crucial decisions that a loon must make while attempting to settle on the best possible breeding lake and rear as many offspring as possible. The Mystery Female, Orange over Mint-burgundy — OMb for short — faced such a decision. OMb returned to the study area as a 4-year-old in 2009 and began hunting for a place to breed. She settled on Upper Kaubashine (at the end of the long arrow below) in 2012 and lost clutches of eggs to predators there in both 2012 and 2013. OMb then

N Nokomis to Bear w UK also

faced a stark choice — remain on a poor territory attempting to breed or abandon Upper K and try to move to a more promising locale.

Shifting from a poor territory to a good one might seem like an easy call to make, but female loons must cope with a shortage of potential mates owing to early senescence and frequent fatal fighting in males. Every year we see many loner females, some of which live on good breeding lakes, waiting for a mate. So it is an open question whether a female should desert a mate and breeding territory — even a poor one — to try and move to a better location. You see, in trying to secure a new territory, a female must temporarily leave her current one, risking its loss to another female on the prowl.

OMb decided to abandon Upper Kaubashine in 2013, establishing herself as the new breeding female on Bear Lake (shorter red arrow), whose female had died. This appeared to be a wise move; Bear had yielded chicks in 9 of the previous 13 seasons, while Upper K had not fledged a chick in 35 years.

Chance plays a big role breeding success of loons, as in all animals. In a curious twist of fortune, Upper Kaubashine stunned lake residents by hatching two chicks from a terribly exposed nest site. Since Bear Lake only produced a single chick this year, OMb’s choice of Bear over Upper K looks like a poor one, as of now. But chick production in these two lakes will probably return to form. If so, and if OMb can hold onto her new territory, her decision to leave a perennial failure for a proven chick-producer will have been a good one.

I must confess that I had mixed feelings to learn this week from Nelson that the North Nokomis pair had hatched two chicks. Despite having seen scores of newly hatched chicks, I still enjoy watching the tiny fuzzballs bob up and down next to their huge parents while the adults, in turn, move gingerly around their tiny young to avoid injuring or drowning them.

So it was, in one respect, nice to learn that the North Nokomis pair had hatched the eggs from the conspicuous nest I had inspected on 25 July, after following the simple, clear instructions from my research team. But I recall thinking, “Oh geez!” on that date, because only two other breeding pairs of the 120 that we follow were still sitting on eggs. (Both of those, we had determined, were cases of infertile eggs that the pair had incubated for six weeks or more.) I have always presumed that chicks hatching in late July or August have too little time to mature physically, develop foraging skills, and learn to fly in time to make the fall migration.

Now we have the data to look at this question robustly. In other words, having captured and marked 983 chicks since 1991, we can determine whether hatching date is a predictor of survival to adulthood. Logically, there must come a date in late summer beyond which chicks run out of time. However, pairs might face a gradual decline in the likelihood of their chicks surviving migration, or there might be a rather sudden threshold date past which chicks that hatch cannot survive.

If we look at all chicks banded from 1991 on, and calculate how many have made it back to the study area as adults, we learn what the figure

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shows. In short, hatching date does not appear to influence survival to adulthood strongly. At the very least, we can say that chicks hatching in mid-July survive at a rate no lower than those that hatch a month earlier. There is a hint of a decrease in survival from early to late hatches, but it is only a hint.

As usual, our data are not perfect. In fact, we have too few cases of very late hatches to gauge the likelihood of the two North Nokomis fuzzballs (which hatched on about 28 July) making it off the lake this November. On the plus side, 470-acre North Nokomis Lake has one of the highest rates of survival to adulthood in the study area. (The territorial males on Gilmore and Cunard were hatched there.) I would like to think that the gutsy North Nokomis pair will be rewarded this fall with two healthy fledglings. So I am keeping fingers crossed for them.

A few months ago I wrote a post about the impact of black flies on nesting of loons. Some might recall that, after abandoning their first nesting attempt, pairs sometimes reuse the nest, leaving the two original eggs in place. This situation produces supernumerary eggs: two addled ones, two still alive. Despite some odd-looking clutches, though, the impact on reproduction seems minimal. That is, the presence of extra eggs in a new nest does not appear to impair incubation of live eggs. Chicks still hatch normally.

In fact, I had all but forgotten about black flies by the time it came to the loon capture season this year. You see, capture is an inherently cheery process. First of all, capture is only possible on lakes with chicks, so we only visit such lakes. Second we work at night and become so absorbed in the demands of creeping up on protective adults and their awkward, fuzzy offspring that the travails of the population at large do not enter our sleep-deprived brains. Between the adrenaline rush following a challenging capture and the warmth of feeling that accompanies the release of parents and their adorable young, nothing else matters.

One issue nagged me even during capture this year though. The great majority of chick broods were singleton chicks (like the one on Muskellunge Lake in Linda’s photo). So few two-chick broods did we encounter that each one seemed an oddity — an almost inconceivable reproductive bounty. 2017 was a surprise, because, based on many previous years of capture, I had come to expect roughly equal numbers of two-chick and one-chick broods.

In the days following my nocturnal boating adventures, I mulled over the abundance of singletons in 2017. It was then when black flies entered my mind. Was it possible that black flies had disrupted incubation to such a degree that many pairs had lost one of their two embryos early and hatched only one chick? This might happen if fly-bitten pairs spent enough time off of their nests that one, but not both, of their eggs became inviable. If so, years with many nest abandonments owing to black flies should also be those with many singleton chicks. In fact, this is the case, as the figure below shows.

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Thus, it seems that black flies inflict a double whammy: they cause widespread abandonment of nests, and nests not abandoned suffer from reduced hatching rate. To make matters worse, cold spring weather, which prolongs the lives of black flies, also causes hypothermia of loon embryos, endangering their survival.

Now I have somewhat simplified the factors that cause singleton chicks in loons. I certainly have to explore additional factors, looking, for example, to see if loons are more prone to laying one-egg clutches during severe black fly outbreaks (although a quick check of the data revealed no such pattern). But it seems that we have yet one more reason to hope for rapid and sustained spring warmup in the Northwoods.

It is July and time to hide the chicks! That’s right; while human parents show off their progeny — perhaps partly to solicit help in caring for them — loons do the opposite. You see, intruders looking to evict territorial residents scour lakes for chicks, because the presence of chicks indicates that the lake contains good nesting habitat and abundant food. So by producing young, a breeding pair has put a giant target on their backs, providing an incentive for any intruder that discovers the chicks (like one of the six intruders shown in Linda’s photo) to return the following year and make an eviction attempt. We should expect, therefore, that parents would hide their chicks from intruders whenever possible.

Of course, breeding pairs are fighting a losing battle. On the one hand, they must feed and protect their chicks, which includes vocalizing often to warn their mate and chicks of passing eagles and other dangers. On the other hand, when intruders fly over or land, parents need to ignore the chicks altogether. Toggling between these two behavioral modes is no small task. Furthermore, while it is desirable to protect your long-term ownership of the territory by hiding your chicks from intruders, you do not want to lose them in the process!

Although chick-hiding is a tricky business, loon families do have a strategy for coping with the sudden appearance of intruders overhead, which fly over at a speed of about 70 miles per hour. We call it “dive and scatter”. At the appearance of a flying intruder in the distance, a loon pair and their chicks quickly slip under water. The chicks swim toward shore and, once there, are hidden by their brown plumage, which makes them resemble rocks or logs. Meanwhile parents swim under water to the middle of the lake, which draws the intruders to them and not the chicks. The aim of this coordinated behavior pattern by chicks and their parents seems clear: keep intruders from seeing the chicks. On its face, dive and scatter behavior clearly seems a means of helping parents’ maintain possession of their territory.

I need to pause here for a second to consider an alternative explanation for dive and scatter. In fact, the most obvious reason why a pair and chicks would dive and scatter is to protect the chicks themselves. Intruders do kill chicks commonly, so this is a viable hypothesis at first blush. But chicks are most vulnerable to being killed by intruders in their first two weeks, so dive and scatter as chick defense — if it is a viable explanation — should occur mainly among small chicks. Yet dive and scatter occurs rarely in small chicks and very commonly in those four weeks and older. So the hypothesis that dive and scatter is a behavior to protect small chicks from intruder attacks can be easily rejected by its timing.

We have known about dive and scatter behavior for some years, but yesterday on Woodcock Lake I learned that loon parents know when to call off the ruse. While feeding their single chick along the lake shore, the Woodcock pair spotted two intruders in flight. The family dove and scattered, the chick hiding near shore and parents making for the lake’s center, in stereotyped fashion. Following the script, the two intruders landed by the parents (and far from the chick), the four adults circling and diving together for several minutes. The charade abruptly fell apart when an eagle flew over the part of the lake where the chick was hiding. Both parents immediately ceased interacting with the intruders, wheeled towards the eagle, and wailed desperately for several minutes, while swimming in that direction. In a half-second, the breeding pair had morphed from cool, detached individuals with nothing to hide into into frantic worry-warts!

Some might view such a loss of composure by a breeding pair to be quite costly. If intruders are able to learn about the presence of chicks by detecting chick defense behavior such as that shown by the Woodcock pair, then the pair exposed themselves to the threat of future eviction by wailing to defend their chick in the presence of two intruders. A clear blunder….until you consider that the alternative was to lose the priceless product of their summer’s breeding efforts.

 

Though it sounds odd to say, people put loons on a pedestal. Their love for loons causes many people to want them to be a bit better than us. Of course, this is the origin of the popular myth that loons mate for life. Such a handsome animal, it seems, should have behavior to match. Besides, loons certainly appear to work so well together in foraging, defending their territory, and rearing young that one can easily imagine that male and female are committed to each other deeply.

Having studied several other species of animals before turning to loons, I did not share the expectation that loons would be good role models for humans. I had seen too many cases of nature red in tooth and claw. In fact, I suppose I expected to learn ways in which loons might not fit the mold of traditional monogamy. What has surprised me the most of all my findings is the degree to which loons have broken that mold. Rather than aiming to remain paired with a single mate throughout their lives, loons seem pursue a far different but straightforward goal. They seek to produce as many chicks as possible by remaining on a breeding territory whenever possible. This strategy requires them to turn a blind eye when their mate is evicted by another loon. Their allegiance is to the breeding territory, not the mate.

No territory illustrates the cold pragmatism of loons better than Blue Lake-Southeast. Recent years have been turbulent on Blue-Southeast. In 2015, the male was evicted by an unmarked male after producing two chicks with his mate. The displaced male hung around on the lake and later regained his position, but last year was a repeat performance; this time a four-year-old male from Bolger Lake evicted the long-term male after hatching and killed the single chick. In a scene worthy of Greek mythology, the Bolger bird actually picked up the lifeless two-week-old chick while chasing its mother across the water’s surface.

Now, if a human mother had witnessed such a grisly spectacle, I doubt if she would have been able to forgive and forget. But loons are not humans; the female whose chick had been killed by the usurper quickly paired with him and remained so this spring. The unlikely pair weathered the black fly emergence in May, hatched a chick in early June, and are now raising that chick on the east lobe of Blue Lake. As the triptych of photos above shows, I caught up with this very tame family yesterday afternoon. At the time, “Chick-Killer” (as my field team affectionately calls the male) was enthusiastically diving for food for the thriving chick, while the female looked on. The dutiful, coordinated parenting of the two adults suggested that they constitute an indivisible unit — that their pair bond would withstand the test of time. But looks are deceiving.

My world gets turned topsy-turvy each year when the first chick hatches. Each year at this time most pairs are struggling just to hold their territories, find safe nesting sites, lay eggs, and fight off the black flies that chew on them. Today, for example, I visited four established pairs that had built nests and laid eggs, but only one — I am proud to say it is the pair on our home lake, Currie — is still incubating the eggs. The others had abandoned their nests owing to black fly infestation. Four lakes is too few to draw conclusions, but my findings today illustrate the debacle produced by black flies again this year. I am just not quite sure how to reconcile the carnage I saw today on my circuit of lakes with the engaging photo that Linda Grenzer took of the Jersey City male alertly guarding his chick.

I suppose the clearest message sent by the news of a successful hatch is that the world is varied and bright spots remain. Indeed, many loon pairs are beating the black flies and thriving. In fact, a few dozen pairs will follow JCF’s lead and hatch young in the next week or two. Moreover, most pairs that have fallen victim to black flies will lay a second clutch of eggs shortly; there is still ample time to hatch and rear young that can migrate south in November. So do not worry if the loons on your lake are swimming distractedly about right now instead of sitting on eggs. They will likely get back to it soon. In the meantime, take heart — as I am trying to — in the photo above.

 

 

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Most of us think of adult females as the main care-givers and protectors of the offspring. I am reminded of this constantly during my work. Several times a year, when I chat with a lake resident about their loons, I hear them say “She was on the nest today” or “The mom was feeding them down at the south end” or “She hollered like crazy when the eagle flew near the chicks”. It is natural for humans to infer that the female takes the lead in breeding activities; after all, we are mammals. In almost all mammals, females protect the young within their bodies for many weeks or months before they are born and continue to care for the young by themselves after they are born. Lengthy gestation makes male parental assistance superfluous, so mating systems have evolved in which males occupy themselves in other ways. Most mammals are either polygynous (one male mated with multiple females) or promiscuous (rapid mating and the lack of a pair bond). Females, in such cases, are left to provide most or all of the parental care, because males are either engaged in seeking out new mates or are not around at all.

Birds are different. Laying of eggs “frees” females, in the sense that they are not physically connected to the eggs and are no better equipped to care for eggs or young than are males. In birds, therefore, monogamy and biparental care are the rule. Most male birds  mate with a single female and provide substantial assistance to her in rearing the young. Male loons, as I have noted in the past, incubate the eggs equally with females and actually provide somewhat more parental care for chicks. Thus, more often than not, when folks tell me that the female hollered at a menacing eagle, or the female was feeding the chick lots of minnows this morning, they have mistaken the male for the female. (This puts me in the awkward position of either correcting the mistake, at the risk of embarrassing my friend, or grinning good-naturedly and leaving the error unchallenged.)

I got one more reminder of male loons’ central role in reproduction during a statistical analysis this past week. I asked whether males or females show age-related changes in fledgling production as they mature from young territory holders to middle-aged to senescent adults. (Note that one bar is missing for each sex in the figure below: too few females settle by age 5 and too few males survive to age 24 to produce reliable estimates of reproduction for those age-classes.) Both sexes show an increase in fledgling production after their first few years on territory, a pattern seen in many animals. Females showsenescence-paper-figure-3

“reproductive senescence”, another widespread pattern, in that fledgling production declines near the end of life. What is surprising here is that males do not show a decline in fledgling production as they reach old age.

What is odd about the male pattern? We have growing evidence that male loons decline at a younger age and more severely than females do. Recent analyses have shown that males lose body weight as they age, and that males suffer a high rate of mortality, especially as they reach their late teens. And, of course, males engage in dangerous battles. So the capacity of old males to produce lots of chicks into their 20s runs counter to what we would expect based on male survival and body condition. How do they do it? We are still puzzling over the pattern, but the most likely explanation is that old males invest heavily in chick production — perhaps through extra feedings of chicks or an extended period of care — to crank out a few more chicks before the wheels come off completely. Hence, old male loons appear to make a “terminal investment” in breeding success. Of course, nothing is free. Terminal investment is a deal with the devil; high chick-rearing success comes at the cost of earlier death.

 

LMG_2782 Muskellunge Chicks Hide from Intruders

Linda Grenzer’s striking photo from two weeks ago got me thinking about loon breeding success. The picture is a sight that will please loon fanciers — two big healthy 9-week-olds resting side by side while their parents circle with intruders. (The chicks are holding their legs out of water, as resting loons commonly do.) Since territorial pairs almost never lay more than two eggs or hatch more than two young, the photo depicts a monster year for the breeders on Muskellunge Lake. Despite black flies and raccoons (which threaten nests), eagles, muskies and snapping turtles (which attack chicks), and constant intrusions by competing adults (which seek to drive adult breeders off of their lakes), the male and female on this lake reared two chicks to adulthood. Quite an achievement!

For Muskellunge, 2016 marks the sixth year in a row of chick production. Ten chicks in all have been raised by the pair during this stretch (including one rehabbed chick we added to their singleton brood in 2014). While I am thrilled to see such an abundance of chicks come from a single lake, it is not the norm. As residents on most lakes well know, loon breeding is a dicey proposition.

Consider, for example, South Two Lake, a normally productive breeding lake where the sudden disappearance of two successful breeders after 2015 left the lake wholly without a pair in May and June of this year, until it was finally resettled by a male and female in July — far too late for nesting. Or look at the Boom Lake-Hodag Park territory, where the pair had fledged five chicks across the past four seasons until the male became entangled in fishing line in the spring of 2016 near Panama City, Florida and never reached Wisconsin. Baker Lake was a consistent chick producer until 2013, when a five year-old male settled there with an unmarked female; they have lost nests to predators each year since then. Most spectacularly, the productive pair on Blue Lake-Southeast weathered the storm of parasites, egg robbers and chick predators only to see the male lose territorial ownership to a young usurper, which resulted in the chick’s death. Finally, pity the pair on tiny Liege Lake or Wind Pudding-West where, despite successful territory defense and incubation, the parents were unable to locate enough food to raise a single chick past six weeks of age.

In light of the many hazards facing loon pairs, it seems remarkable that we ever see a photo such as the one above. As one might surmise from the preceding paragraphs, there are several requirements for successful breeding. Abundant food is essential, of course. Each year, many pairs attempt to breed on tiny lakes where food limitation prevents them from rearing even a single chick, let alone two. Nesting habitat is vital; lack of boggy or marshy shoreline or an island prevents many pairs from even attempting to breed. Two less obvious factors can make or break a breeding effort: 1) the ability of both pair members to maintain their breeding positions throughout the season despite an onslaught of young adults looking to evict them, and 2) the male’s familiarity with proven successful nest sites, which dictates whether the nest is positioned in a location likely to survive four weeks of incubation.

Some breeding pairs have everything going for them, so that they surmount all obstacles and raise chicks to fledging year after year. Muskellunge (see photo) is large enough — at 160 acres — that food abounds, and the lake features several shoreline zones that support nesting. The resident female is much larger than average and aggressively repels intruders that challenge her. She is paired with a tough 18 year-old male — a bird experienced enough to know the locations of multiple nest sites on the lake but young enough to be in good physical condition and not vulnerable to eviction. In short, Muskellunge Lake is currently in a “sweet spot” for raising chicks, like Townline, Manson, Little Bearskin, East Horsehead, and Buck Lake. Lakes that get on a roll like these produce a disproportionate number of chicks, which will mature, return, and sustain the population. In time, these productive territories will falter, owing chiefly to the loss of one or both members of the vigorous, experienced breeding pair. For now, though, let’s enjoy the bounty of offspring that these lakes produce and look forward to the emergence of new productive territories that will succeed them.

 

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.