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!

 

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.

 

2016-08-01 02.16.10

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.

2016-07-31 01.32.29

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

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

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

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

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

The goose population has been increasing in northern Wisconsin. Ten years ago, a Canada Goose was an infrequent sight on one of our study lakes. Geese were confined chiefly to a few large lakes and seen overhead only as they migrated northwards. But no more. Now we encounter adult geese and often pairs with goslings on about half of our study lakes.

Apparently the increased availability of waste grain from agriculture, the proliferation of lawns, and increased temperatures have fueled the explosion in goose numbers, which has the potential to impact loon reproductive success. You see, geese and loons both prefer nesting sites safe from mammalian egg predators like raccoons and foxes. Such sites are often on islands and are limited in number. If a small island offers the only available safe nesting on a lake, goose and loon pairs are both likely gravitate to it.

The problem is not merely that loons must now compete with geese for nesting sites; geese actually get “first dibs”. While both loons and geese incubate their clutches for about four weeks, geese initiate their clutches two weeks or so in advance of loons. (The earlier onset of goose nesting is evident right now in our study area, as many goose pairs are rearing their broods of two to six goslings, while the earliest loon nests will not hatch for another week or ten days.) On occasion geese and loons nest within a few meters of each other on islands — as we observed on Oneida Lake a few years ago. Coexistence between the species is possible. But the presence of a sitting goose appears to discourage loons from nesting nearby, which often forces loon pairs to select sub-optimal sites for incubation.

Consider the plight of the Clear Lake loon pair. The safest, most desirable nest location on Clear Lake is a small shrubby island off of the public boat landing. Loons have shown a strong preference to nest on this island during most years. This year, the geese got their first, and the loons had to settle for a new nesting location on a long peninsula about two kilometers south. They may hatch chicks off of this peninsula, but the site is not offshore, like the shrubby island, so it is clearly vulnerable to mammalian egg predators.

I am optimistic that this spunky loon pair will be able to pull off a hatch. As can be seen from our shaky video, the male (on the right in the video) is still working on improving the surrounding nesting area, and the pair has a well-constructed nest with two eggs (visible to the left of the female, after the camera pans left). Still, this well-constructed nest will not protect them from a scavenging raccoon that ambles by.

 

 

LMG_5401 Crystal Lake NestOn its face, the photo is comical. A loon sits on its egg on a swim raft, a meter from an American flag. A garish McMansion is visible on the opposite shore. Few photos of the common loon are more at odds with the image of the species as a symbol of the northern wilderness. Yet Linda Grenzer’s photo of the nest on Crystal Lake is a poignant portrait of the situation faced by most territorial pairs in north-central Wisconsin.

Many popular accounts describe common loon populations as holding on for dear life. To be sure, loons are threatened by shoreline development, which reduces nesting habitat. Recreational fishing and boating kills many adults and chicks prematurely each year. And methylmercury in the food chain, swollen by coal-based power production, likely impacts the loon population as well.

Against all odds, however, loons appear to be holding their own in northern Wisconsin. Despite consistent losses caused by collisions with boats and entanglement in fishing lines, populations have bounced back in recent decades. The surprising resilience of loons seems to result from two facts. First, loons prefer to nest on small islands and on boggy and marshy habitats that humans avoid. Thus, loon pairs continue to produce young on lakes that are virtually encircled by human dwellings. Second, as illustrated in the photo, loons tolerate — and rarely even benefit from — human alterations of aquatic habitats.

The pair on Crystal Lake, whose nest is pictured, were in a real bind. Crystal is a very pleasant lake, but it lacks islands — the nesting spots most favored by loons — and also features neither bog nor marsh. Previous nesting attempts by the male (reared on nearby Muskellunge Lake in 2006) and a parade of 3 or more different females since 2011 ended in abject failure, the eggs an annual donation to local raccoons. Although they crawl onto it with great difficulty and complete loss of dignity, the Crystal pair decided a few weeks ago to place their nest on a low-lying swim raft. The raft is high above lake level and not equipped with any sort of ramp to assist small chicks in re-mounting the raft once they have entered the water, so we are concerned that the chicks will not be able to return to the nest (to be kept warm by a parent in the first few days of life) once they have left it. Still, the likelihood of hatch is good, and Linda and her helpers placed nesting material on the raft to keep the eggs from rolling off. We are hopeful that the loons can cope with the problem of nest height just as they do with a host of other anthropogenic obstacles each day in northern Wisconsin.

I shall keep you informed. If the Crystal pair can hatch and rear their chick to adulthood, they will be a vivid example of the capacity of loons to adapt and thrive in an environment thoroughly dominated by another species.

LMG_4295 one day old chick on back

After 2014’s disastrously low chick production owing to black fly infestation, I was anxious that breeding success would rebound in 2015. My concern defied reason. Loons are long-lived, of course, and a few or even several consecutive years of below-average chick production are not enough to have a lasting negative impact on the population. Still, blinded by the irrational human tendency to infer long-term impacts from short-term patterns, I wished to see our loon pairs come back and raise many chicks in 2015, just as Linda Grenzer’s pair has (see photo).

This year is shaping up to be a very good year indeed. As of this date, 31 of the 120 territories we follow (25.8%) have already hatched chicks. In contrast, only 2 breeding pairs (of 112 pairs; 1.8%) had chicks on this date in 2014. That is an unfair comparison, because the black flies devastated early nests in 2014. However, 2013 was a rather good year for chick production, and in 2013 only 20 of 108 pairs we followed (18.5%) had chicks on this date. Although these data are slightly biased because of yearly differences in nesting schedules — that is, 2013 was a slightly later year than this year — 2015 should be an awfully productive breeding season for loons in northern Wisconsin. As a scientist, I am breathing a huge, sheepish sigh of relief!

LMG_3382 JCF Male

We have almost completed the first round of visits to study lakes. Our ever-expanding list of lakes makes this no mean task, as our list has grown from 95 territories in 2008 to 123 this year. Kudos to the field staff of Joel, Eric, Chris, and Linda (whose photo appears above), who have worked hard to visit lakes and ID loons from leg bands — and also Al on West Horsehead and Pat on Silver Lake, who have e-mailed with data from their lakes.

Of 162 banded loons on territories as of late 2014, 141 (or 87%) have reclaimed those territories this spring. Among 21 missing territory holders from 2014, 4 have been resighted in 2015, but were apparently evicted from their territories early this year. Thus, the minimum survival rate from 2014 to 2015 for territory holders is about 90%. This figure agrees closely with survival rates calculated from Wisconsin and elsewhere within the breeding range.

Sky-high return and survival rates for our population highlight a simple pattern in the life history of common loons. In ecological terms, loons are “K-selected”. That is, they are long-lived, take several years to reach sexual maturity, produce few offspring during their lives, and invest heavily in parental care for the few offspring they do produce. Loons are not explosive reproducers that “shot-gun” many offspring out into an uncaring environment in hopes that a few survive. Rather, they work hard to maintain ownership of their breeding territory, eke out one or two chicks a year (in a good year!), and defend their chicks vigorously against all comers!

By any reasonable measure, 2014 has been a dreadful year for loon reproduction. Even if we capture every remaining loon chick that we know of, our total number banded will be almost exactly half of last year’s total. Clearly, the black flies hit the loons’ breeding efforts in the mouth, and they could not recover. Still, it is only one year. As my work has shown me over the past two decades, breeding success is a roller-coaster. In fact, if you combine last year’s bumper crop of chicks with the withered output of this year, the message is that loons are producing enough chicks to sustain the population. So all is not lost.

There have been a few bright spots this year. Though we were alarmed in early June when the Jersey City Flowage male ingested a fishing lure and became hopelessly entangled, the folks at REGI were able to save the bird, he was released near his old territory and recovered — except for possession of his territory! The REGI folks were brought an emaciated chick found by a roadside that they fed back to reasonable health and that we were able to foster successfully to a loon family that has raised it as their own. We recently banded both this fostered chick and its sibling, a true biological chick of the pair. Both chicks have become big healthy, strapping young birds, thanks to their parents’ tireless efforts.

In short, life goes on. I am confident that we will long remember 2014 and not wish to repeat the experience. But perhaps we should best remember 2014 as a year that, while dismal for chick production, was balanced out by strong reproductive years surrounding it and did not pass by without a few cheerful tidings.