Juvenile loons are in a race against time. While their parents seem to relax following the breeding season — wandering from lake to lake as if on a goodwill tour — juveniles, like the three-month-old in Linda Grenzer’s photo, face a ticking clock. After hatching in June or July, juvies must reach near-adult size by ten weeks of age, practice takeoffs and landings, and become strong enough to make flights of hundreds of miles on their southward migration in early November.

They are racing the ice. Temperatures cool in September, become unpleasantly chilly in October, and truly plummet in November — and lake temperatures follow suit. Ice-up can occur anytime between mid-November and mid-December in northern Wisconsin, and ice-up is the end of the line for juveniles. Opportunistic bald eagles await juveniles that are not prepared to migrate and become trapped in the ice. Apparently sensing the desperate task that will confront their offspring in the fall, parents stuff them with fish for eight long weeks in July and August. Chicks grow explosively during mid-summer. But they face their most challenging task in autumn, when parental support wanes and they must learn to feed themselves, improve their body condition, and prepare for their southward journey.

In general, scientists have paid little attention to the juvenile period in birds. Our neglect is natural enough. The breeding season is chock full of interesting behavioral and ecological events: pairing of mates, defense of breeding territories, selection of nest sites, and relentless territorial intrusions by nonbreeding adults seeking to settle. Perhaps ecologists can be forgiven for focusing their attention on breeding behavior and trusting that juveniles will take care of themselves.

But we wondered. If young adults settle on breeding lakes that closely resemble their natal lakes, might juveniles — which must fight for their lives just to become adults — also exhibit clear preferences for certain kinds of lakes over others? Constrained by flightlessness to forage only within the lake where they hatched, we might expect juveniles to become highly specialized to hunt and consume the species of prey found on the natal lake. So once they become capable of flight, we might expect them to visit and forage on other lakes very similar to their natal one. That is, juveniles reared on a diet of bluegill sunfish and used to hunting that species should spend most of the pre-migratory period visiting lakes full of bluegill that they can catch and consume efficiently. And juveniles accustomed to eating snails and leeches should find lakes full of those invertebrates on which they can feast.

Our interest in lake visitation patterns of juveniles during fall inspired us to plot the local movements of youngsters between lakes in the fall of 2012, 2013, and 2014. Kristin, Gabby, and Nathan used their band-spotting skills to locate juvies in September and October of these years. They found close to 200 cases where a juvenile we had marked had flown to forage on a lake other than its own. Using these data, Brian, who joined us this summer, asked, “Do juveniles forage on lakes at random, or do they prefer to forage on lakes like the one that hatched them?”. As the figure below shows, the mean difference in pH between a juvenile’s natal lake and the lake where we spotted it foraging (red vertical line) was far less than the distribution of differences we would have expected, if juvies had foraged randomly (grey bell-shaped curve).

Z_pH_Randomization

Although Brian has a few statistical checks to complete, the pattern seems clear. Juveniles exhibit strong preference for lakes that resemble their natal one in two respects: 1) pH and 2) water clarity (data not shown). Brian’s analysis is ongoing, and he is trying to learn how closely these chemical and physical attributes predict the food available to loons in a lake. But we are betting that the stark preference of three-month-old juveniles for lakes that remind them of home occurs for a simple reason. Juvies try to spend their time hunting prey in familiar conditions to build themselves up for their most dangerous first southward journey.

Last year I wrote a blog post in which I concluded that late-hatching chicks returned at a rate no different from early-hatching chicks. I found the result surprising, as one would expect early hatchlings to have a head start in learning to feed themselves, honing their flight skills, and preparing for their first migratory journey. The photo and story I got from Linda Grenzer a few days ago has forced me to wonder if I need to collect more data on this question.

The breeding pair on Squaw Lake had an eventful year in 2018. Delayed, like all other pairs, by the late thaw, they initially nested along the shoreline near the boat landing. After a predator snatched both eggs off of the nest, they nested again not far away. This time they were more fortunate; the eggs hatched, but not until about July 22. When we captured the family on August 3rd, we found the chicks almost comically small — two little puffballs that did not approach the size of the many other juveniles we had encountered. Chicks are cute in their first few weeks, and we enjoyed observing them and handling them cautiously while giving the female a new set of bands.

Our delight at seeing the adorable chicks was tempered by the fear that chicks hatched so late would not mature in time to complete the southward migration. The fear is justified; parents must balance the energetic demands of their demanding offspring against their own need to maintain good body condition and prep for their autumn journey. Inevitably, adult loons spend progressively less time on their home lake in September as they forage intensively, molt into drab winter plumage, build up fat levels, and, in late October or early November, head south. This goes for parents and non-parents alike.

So it was not surprising to get a report from Linda that the Squaw adults had left their breeding lake, leaving their late-hatched chicks to fend for themselves. What was alarming was that one chick had chased someone’s jig, managed to hook itself above the base of the bill, and was no longer diving or foraging normally. Further evidence of its desperate condition was that it was not difficult to capture and weighed a mere 1750 grams — roughly 1 kg less than it should have at 9 weeks. Following an X-ray at Raptor Education Group, Inc. in Antigo, the chick was found to have swallowed a second hook from a separate encounter with an angler.

Since we have long since ceased our routine visits to study lakes, we can only speculate about the series of events that put the chick in this bind. Marge Gibson of REGI suspects that, without parents to help it satisfy its foraging needs, the chick was struggling to feed itself. In its desperation, the chick began to attack fishing lures until the hook in its cheek and weakness conspired to incapacitate it.

If Marge is right, and late-hatched chicks are sometimes left with too little feeding capacity to maintain themselves, then this pattern should show up in our data. Specifically, we should see fewer very-late-hatched chicks return as adults to the study area. This plausible scenario will fuel another round of data analysis…when I find time!

To end on a positive note, the angling victim is bouncing back at REGI and feeding voraciously. If you do not believe me, look at this video from the REGI website.

https://www.facebook.com/RaptorEducationGroupInc/videos/470615703434171/

If it continues to thrive, the REGI folks will face another challenge: what to do with a healthy juvenile, but one whose stay in captivity and recovery made flight practice impossible.

 

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

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

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

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

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

1) the adjacent Nokomis-East Central territory,

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

3) Soo Lake, 11 miles to the NE,

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

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

6) Miller Lake, 18 miles N,

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

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

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

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

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

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

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

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

 

As many of you know, I am a worry-wart. Normally I get so stressed-out about my kids, my teaching, my research, my health — and many other matters that are going well — that I hardly have time to obsess about loons in the study area. But Linda Grenzer’s bleak photo of conditions on her lake today gave me a jolt. Could the late ice-out that we are facing in 2018 delay the season so much that it damages the breeding prospects of our loon population?

One might think that the later the ice comes off of the lakes, the later the loons nest, and the less time parents have to fatten up their chicks and prepare them for their first southward migration. Thus, a late ice-out might well lead to reduced breeding success for the population. Although there are many “if”s in this string of logic (and a preliminary analysis did not bear out the pattern), I felt concern  gnaw at me.

So I did what scientists often do to stave off despair: I looked at the data. First, I looked to see if loons nest later when the ice goes out later, which almost has to

Screen Shot 2018-04-15 at 8.46.07 PM

be true. It is true, but there is a lot of noise in the data. That is, loons are constrained to nest somewhat later in years when the ice goes out later, but the picture is not simple. The reason for the noise becomes clear when you look at the lag time between when loons settle on their territories and when they hatch their

Screen Shot 2018-04-15 at 8.43.02 PM

young. There is a very strong pattern here. When the ice goes out early (left side of graph), loons dawdle and wait weeks before nesting. But when the ice goes out late, as it will this year, pairs get down to business quickly, nesting within a week or so of territory return. So loon pairs are somehow able to catch up in years of late ice-out so that their breeding schedule does not differ greatly from other years. (Notice also that the orange line in the top graph is flatter than the blue line.)

What accounts for this pleasing pattern? We can make a pretty good guess based on findings in other migrant birds. Spring migration is an energetically costly process. In an early year, the ice is gone so quickly that loons settle on their lakes as soon as they return from the wintering grounds. In such cases, their fat levels are very low from migration when they first occupy their territories, and it takes a good deal of foraging before they return to good condition. In a late year, loons cannot settle on their territories right away but must wait on nearby rivers that have open water. There, they are able to forage and restore their bodies to good condition. As a result, loons hit their territories in prime body condition and fully recovered from the migratory flight in years of late ice-out. Thus, they can get down to breeding quickly.

Although I was heartened by the data I saw above, I had a look at the numbers that most directly addressed my concern about late ice-out and population breeding success. There is a no statistical tendency for the population to produce more loon chicks in years of early ice-out, despite the many years of data we have to look for such a pattern. Indeed, some of our best years for loon breeding (2013, for example) have occurred when the ice goes out late. So those many of you shivering in northern Wisconsin and other frigid regions can relax about one thing; the loons are no worse off in years when spring comes late than when it arrives early.

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

Screen Shot 2017-08-12 at 11.24.21 PM

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.

LMG2693 Loon Between the Ice

I always dread ice-out. While I am excited to know that I am scant weeks away from seeing the loons, ice-out now tends to occur during the high-stress period of the Chapman spring semester — and I am seldom ready for the event. As Linda’s photo from the Wisconsin River shows, loons are coming back, accumulating on the river, waiting for their breeding lake to open up enough to permit landing there. Meanwhile, my Animal Behavior students are sweating their behavioral experiments and write-ups.

Let’s focus on the positive of early ice-out for data collection. Rain and warming temperatures mean that ice-out is only days away. Soon we will begin to log the identities of returning veterans anxious to attempt another year of breeding. Since the lakes will be habitable two or more weeks before the historic mean date of ice-out, while many returning birds are still en route from the wintering grounds, we have an opportunity. In theory, adult loons that are in good physical condition generally should be those that can complete the breeding molt early and also migrate early. So the loons that show up first on breeding lakes should be those in good condition. Recent findings have shown us that this group comprises males and females about 8 to 15 years old — the prime of life for a loon. Thus, we predict that the early arrivals are in this age-class. The laggards should be breeders that are either very young — 5 to 7 years old — or very old — 20 years and older. We might expect the territories of such individuals to remain vacant for a week or more after the ice has come off of them.

If age does turnout to be a good predictor of date of territory occupation, then late return from migration could be another source of trouble for an old established breeder. That is, an old territorial bird whose body condition has begun to decline might not only need to worry about being evicted by a young, fit nonbreeder in the midst of breeding; danger might also come from the tardy return of the old bird to its territory in the spring, which could open the door for a youngster to seize the territory, pair with the old bird’s mate, and hold off the former owner when it returns.

I have painted a dire picture. We will have to use the increasingly early ice-outs like this year’s to measure date of return accurately and see if early ice-outs truly destabilize territory ownership. At the moment, I will tantalize you by reporting that breeding success across the population is higher when ice-out comes late. It is speculative at this point, but this pattern might indicate that early ice-outs lead to ousting of old, experienced breeders from their territories, which in turn suppresses chick production of the entire population. If so, I have one more reason to rue early ice-outs!