High Return Rate in 2015

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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!

Gabby’s Paper on Chick Defense Accepted for Publication!

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Sorry to trouble you with posts on back to back days, but we just got good news from the Journal of Avian Biology. A paper by Gabby Jukkala and me that describes chick defense of loon parents towards decoys has just been granted final acceptance. We are delighted, because we have forged our way through numerous revisions of this paper over the past year or so. It is nice to see that our labors were not in vain.

In fact, the struggle to get this manuscript published is a good illustration of how peer review can lead to new perspectives and discoveries. The paper quantifies the defensive responses of parents to a decoy of an adult intruder; intruders attack and sometimes even kill small chicks. Gabby and I had been able to document that parents of small chicks (0 to 2 weeks) remain near them when a decoy is placed nearby (apparently to ward off surprise underwater attacks), whereas parents of older chicks (4 weeks+) confront the artificial intruder. But in response to reviewer comments, we sharpened our analyses and discovered two more behavioral patterns. While we had long known that males are especially apt to yodel when they have small chicks, we learned through this improved analysis that males with TWO chicks are four times as likely to yodel as males defending a SINGLE chick. This find suggests that males increase parental care in response to the value of the chicks. In addition, we noted that males are more than twice as likely to penguin dance in defense of their chicks as females.

In short, our new paper clarifies our picture of chick defense in loons. Males shoulder a disproportionate share of the burden for chick defense, as we document. Males spend far more time with chicks than do females, yodel at intruders (which females cannot do), and penguin dance much more often than females do. Moreover, male behavior is not mindless, all-out aggression. Indeed, it is nuanced, as males’ toughness towards intruders is combined with a cold calculation of how they can best maximize their Darwinian fitness.

Brace for Black Flies!

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black fly

Black flies have a terrible beauty. From a distance, they are merely pests, which we must tolerate for a few weeks each year. Up close, they are carefully crafted machines with specialized sensory capabilities and sharp tools sculpted to pierce the skin of mammals and birds and extract the blood they require to nourish their eggs. You have to respect them from the standpoint of evolutionary design.

Yet I spend little time admiring these creatures. Indeed, I dread the two-week period in early May when adult female black flies (Simulium annulus) emerge in vast numbers and descend upon loons. Black flies render each day a trial for loons, which are trying to locate nest sites, mate, build nests and prepare themselves for four weeks of joint incubation. Hounded by flies throughout daylight hours when they are above water, loons drop their normal routine and begin to dive incessantly to minimize blood lost to the flies. Breeding pairs that have completed nests and laid eggs are often forced to abandon efforts to incubate them owing to the relentless insects.

In the past several days, fly populations have spiked, and loon pairs are now suspending their breeding efforts for the moment and simply ditching the flies. We hope that loons’ nesting efforts are not so closely synchronized with fly emergence this year as they were last, when black flies reduced loon breeding success dramatically. If this is a typical year, a small percentage of loon pairs (say, 10-20%) will abandon their eggs on account of the flies, but many more pairs will persevere in spite of the pests. So our hopes for a banner year of chick production are, as yet, intact.

Although their travails pale in comparison to what loons face, even loon researchers are impacted by black flies. Eric, Joel, and Chris are in the final stages of covering the study lakes for the first time. Since they are trying to observe leg bands of loons that are diving constantly to avoid flies, the LP team will confront a steep challenge in the next few weeks. I hope they are up to it!

Jersey City Flowage Male Healthy and Back on Territory!

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We were horrified last spring, when one of our best-known territorial males — and a super tame bird to boot — became hopelessly entangled in fishing line. This male, the long-term resident on Jersey City Flowage (“JCF”) had swallowed two lead sinkers, and monofilament line was wrapped tightly about his bill, making it impossible for him to eat. In short, the JCF male was destined to perish quickly from lead poisoning and/or lack of food. However, the folks from REGI freed him and used new technology to pluck both lead sinkers from his gut. Although he lost much of his tongue because of the fishing line, we were heartened when Linda found him in July of last year gorging himself on bullheads near his old territory (see photo below). LMG_1395 Jersey Male n Bullhead

But recovery after a brush with death was just the beginning for this bird. In order to reclaim his previous position, the male needed to regain his strength, migrate successfully to his winter quarters, survive the winter, migrate back north, and then — most challenging of all — regain possession of his former territory, which had been quickly snapped up by another male after the injury. (The new male bred successfully last year and was banded by our team.) Yesterday, Linda confirmed that the old, war-torn JCF male has negotiated all of these hurdles and settled in a breeding pair back on his old territory (see Linda’s photo below). We presume that he is also with his old mate, but she is one of the few wily adults that has managed to elude us, despite several efforts at capture. So she is still unbanded. LMG_3377 JCF Male The resilience of the JCF male is important. He is only the third adult to have been severely injured by fishing tackle, lost his territory, and recovered to reclaim it. Three data points can never be a robust sample, but we also have many cases where entanglement occurred and adult loons did not recover. In fact, we have about 9 well-documented cases of death following entanglement, which, combined with the three success stories, yields an estimate of 25% likelihood of recovery. So while the story of the JCF male’s journey from disaster to full recovery lifts our spirits, it is hard to forget that this is the exception, not the rule.

Spring Returns…and Loons Scramble

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LMG_3148 SOO ABJ It is easy for me to say, I suppose, because I am sitting here in southern California in my shorts and t-shirt, wondering only if we have enough lemonade to survive the day (and enough water to make lemonade)! Still, I think most of us can agree that the season is beginning to turn. This week’s highs in Oneida County will be in the 60s, which should take care of most or all of the remaining ice on the lakes, especially with the help of the wind. After a rather brutal winter, we have an ice-out that is about a week earlier than average. Early iceout created an odd spectacle on many lakes this week past: open water devoid of loons. To be sure, breeding pairs are trickling back. Joel Flory has confirmed that both members of the breeding pair on Manson Lake have returned. Lake residents have spotted a pair on Lake Mildred and one of two pair members on Sherry. Linda Grenzer reports that “Clune”, the male on Muskellunge Lake, returned on Friday for the first time, although his long-time mate, “Honey”, has not shown up, and he is currently frolicking with a new female (see Linda’s photo, above) that we banded as a chick in 2004 on Soo Lake, Linda reports. (We are not judging!) Why would territory owners leave their lakes undefended, especially at a time when many adult loons without territories are on the prowl, anxious to seize any vacant lake? The answer is simple. Weather changes rapidly. As migrants that must fly hundreds of miles between the wintering and breeding grounds, loons face a meteorological puzzle. If they molt their feathers and migrate too early to the breeding grounds, they will encounter wintry conditions and uninhabitable frozen lakes on arrival, struggle to find enough food on open water along rivers, and ultimately settle on their breeding lake in poor condition. They will then be at risk for losing their territory to a fitter, stronger usurper who times his or her arrival better and remains in better condition. If, on the other hand, they wait too long to migrate, they might return to find a squatter established on their territory. In such cases, a territory owner would have to battle the squatter to reassert itself as owner. In short, gauging when to return to the lake you own is an inexact business for a territorial loon. We can understand why they might often arrive a bit too early or too late. So we must be sympathetic about the pitfalls of long-term planning and content with a steady trickle of returning loons. Don’t worry. Territorial loons have evolved a sound set of strategies for coping with fluctuating weather conditions — and interlopers. We expect to see most of them re-established on territories within a week. I will keep you posted!

The End of a Journey: Foothold Paper Accepted by Animal Behaviour

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2015-01-31 11.10.42

Publishing papers in scientific journals is hard work. It requires patient and well-planned data collection, thoughtful statistical analysis, and painstaking writing and editing of a manuscript. And then the real work begins! Among the dozens of scientific journals that might publish the paper, one must select a journal that suits the topic of the paper, employs competent editors and reviewers, and makes its published papers available to a wide audience so that it will be read and cited by many colleagues in one’s field. Most crucial, one must convince the journal’s anonymous reviewers that the findings reported in the paper are robust and valuable. In short, publishing a paper is a journey.

Recently, we completed a successful journey, as our paper describing the process of territory settlement in young loons was accepted by Animal Behaviour. Although we would like to celebrate this event, we are more relieved than joyous; relieved because the paper represents a vast amount of field work, number-crunching and writing and became long and unwieldy enough that it earned harsh criticisms from some reviewers. So its publication became, as some journeys do, a story of survival in the face of adversity.

The paper will make available a trove of valuable findings. We report in the paper that young loons do not adhere to the most prominent model for territory settlement. This idea, termed the “foothold model”, maintains that young animals in search of breeding territories target a small set of established territories for intrusions, gradually gain confidence through increased familiarity with that limited set of territories, and then evict the owner of one of those territories (or outcompete another young loon for the vacancy, in the event of the owner’s death) in order to claim the territory as their own.  We show in our paper that, instead of using a foothold of this kind to gain a territory, young loons merely settle on a territory that is similar to their natal one. In some cases, they are able to occupy a vacant territory and breed there. In other cases, they wait to mature and improve in body condition and then evict an owner. But the repeated intrusions that young loons make to specific lakes are not attempts to build upon their familiarity with the lake and thereby increase their competitive ability there.  Rather, they are efforts to assess the fighting ability and perhaps the motivation of the current owner to defend its territory so that the young loon can judge when an attempted eviction is likely to be successful.  Reviewers described our findings as “provocative” within our field, and we hope they are right!

Thanks to all our supporters, especially landowners and friends, who allowed us to study their loons year after year. Publication of this meaty paper is evidence that our mutual investment in loon research is paying off.

Loons Senesce!

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There are three main reasons why I do field research. First, I love the outdoors. While it is unpleasant and inconvenient to many, the field is my briar patch. Second, I love observing and being with animals and gain new insights by watching loons up close and not just looking at data in my office. And third — well, I enjoy looking at data in my office. Complex puzzles and contradictions concerning loon behavioral ecology often become comprehensible when I look hard the numbers and run statistical tests.

Yesterday I had one of those Eureka! moments during data analysis. For the past several months, I have been looking at factors that: cause loons to: 1) be evicted from their territories and 2) disappear from their territories (that is, die). Some months ago I shared with folks that having your mate get evicted puts you at risk for eviction yourself. That is an interesting pattern and one that I continue to explore. But yesterday, I conducted the most sensitive and powerful analysis to date on the potential impact of age on rate of eviction and death in territorial residents. The  results were clear: age has a substantial impact on both eviction and death rates. Specifically, old loons are at risk for being booted off of their territories and they are even more strongly at risk for dying. The pattern was a bit tricky to detect, because very young adults are also at risk for eviction and death. So loons — like humans — have a “prime of life”. Loons of 4 to 6 years of age are still developing and improving in condition; at this age, they have a rather high rate of eviction and death. They reach prime condition about age 7 or 8 and remain in good condition until about 20, then condition falls off again, exposing them anew to a high rate of eviction and death.

The figure above shows the shape of the pattern with respect to age and death rate, when other factors are controlled. Sorry for the arcane title of the Y axis, but here is how to interpret the pattern. The curve shows how often territorial loons die, higher values indicating a higher death rate. The values are higher for young ages and old ages. Thus, the “U” shape shows that young and old loons die at a higher rate than do loons of moderate age. If you look closely, you can see that old age has a much stronger, more consistent impact on mortality than does young age. So senescence is very clear from the graph.

If you are a loon aficionado, you know that male loons often die in the course of defending their territories, while females rarely do so. How does the new finding of senescence fit in with lethal contests among males? On the one hand, it fits, because senescence might make male loons “desperadoes”, who fight hard for their territories because they have no future to lose. On the other hand, the pattern of senescence cannot by itself explain male-only fatal battles, because females and males both senesce. Oh well….I am always looking for another excuse to look at data in my office.

A Last Rendezvous Before Winter

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LMG_3139 Clune n Honey Return

 

As this beautiful photo by Linda Grenzer makes clear, time is passing. Chicks have matured and left their natal lakes, territorial pairs have scattered. Juveniles and adults alike are preparing to head south. Most obvious to loon watchers, adults have shed their striking breeding plumage and donned dowdy winter garb.

Although we might mourn the significant loss of beauty that occurs among loons in the fall, molting is a pragmatic biological pattern. Feathers wear out with time and use, requiring replacement, so seasonal molting of some kind is essential to survival. But why do loons lose the white wing spots, the necklace and chinstrap, and the elegant black head? As usual, the answer is that we cannot be certain without experimental investigation. However, these prominent features of the plumage signal readiness to defend a territory and to breed. Hence, they are inappropriate and possibly costly signals to send during the autumn. An adult that maintained breeding plumage during the fall might attract the attention of others and trigger territorial interactions at a time when it should be feeding vigorously in preparation for migration. Moreover, the contrasting breeding plumage would probably make a loon more conspicuous to predators such as sharks, which could be costly during a long winter on the ocean.

The two loons in Linda’s photo are her breeding male and female — and two of our best-known individuals. They appear often in her photos, such as when 5 intruders came to call and when they refused to abandon their nest, despite merciless black flies. After all they had been through this year, it was a surprise to see the male (Linda calls him “Clune”) and the female (“Honey”) spend a short time together on the brink of fall migration, long after most pairs had dispersed. I find this photo poignant somehow. I suppose it symbolizes to me this pair’s unwavering unified front in the face of all challenges and changes that confront them.

An Inseparable Pair

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With the first snowfall in the offing and adults and chicks feeding voraciously to steel themselves for migration, an odd and touching event has occurred. We have been able to track the local movements of a pair from their breeding lake to other lakes in the study area. This is not a traditional pair of male and female, but a father and his son who have remained together for an unusual stretch. Hatched on July 1st, the Fawn chick was attended closely by his father from the start. When his mother swallowed a fishing lure and anglers cut the line, leaving her trailing yards of monofilament, she abandoned her mate and offspring for several days to recover. Still his dad remained with the chick and increased his feedings to make up for the absence. (The female survived the ordeal and returned to Fawn Lake afterwards to resume parenting.) Dad continued to feed his son assiduously as he grew, matured and began to take flight. Two weeks ago, however, Fawn Lake was empty of loons. The sharp-eyed residents of Lumen Lake, right next door and almost touching Fawn, reported the sudden appearance of an adult loon and chick that turned out to be the missing father and son from Fawn. The two remained on Lumen for a week or so but then disappeared again. Judith Bloom, who for years has monitored several breeding pairs on huge Lake Tomahawk, e-mailed on Thursday to report that she had found (and ID’d!) an adult and chick feeding in a bay near her home that were not from any territory on Tomahawk (see Judith’s photos). Sure enough, the Fawn father and son had made another appearance in foreign waters. BloomTomahawkFawnMandck4 BloomTomahawkFawnMandck6

Now, it does happen that parents continue to feed their chicks at 13+ weeks of age.  What is unusual is that a parent-offspring pair has remained together on not just one shift to a foreign lake, but two shifts. Moreover, the second shift was a whopper, as the duo flew about 6 miles together to land on Lake Tomahawk. While it is tempting to view this event as a reflection of the trend that is becoming routine in human society — offspring remaining with their parents well beyond the normal age of independence — it might warrant scientific scrutiny. In fact, such “fawning” behavior by a father towards his son makes sense evolutionarily, providing Dad has improved his son’s chances of surviving migration and its first winter while still maintaining his own health. In three to five years, when this chick stands to reach adulthood, we will see if the father’s tireless investment paid off.

Chicks Take Wing

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LMG_2826 Three Month Old Chick Wing Span

I just got a report from Kristin, who is still in the study area. The chick hatched and reared on little Buck Lake popped over to Crescent Lake for a visit. Kristin was excited, because she had seen many cases of juveniles flapping their wings and practicing takeoffs, but this was the first flight of a juvenile she had recorded to a nearby lake. Thus begins the phase of juvenile wandering that characterizes the fall months. If this year proves typical, trips of maturing juveniles to neighboring lakes — sometimes up to 10-15 miles from their natal lake — will abound in the next 9 to 10 weeks.

But why do juveniles abandon the comforts and familiarity of their home lake to venture to lakes unknown? This is a difficult question with many possible answers. First, let’s think about the downside of such movements. In deserting its natal lake, a juvenile will be faced with new food sources to which it will have to adjust. Since it must feed itself and mature rapidly to put on weight for fall migration, leaving home seems a gamble. A new lake will also contain new risks in the form of predators or aggressive territory holders (although aggression is generally low in the fall, to be sure).

In order for wandering to new lakes to be a sound behavioral strategy, we would expect there to be advantages that more than offset these risks. One can imagine a number of potential benefits to shifting to a new lake. In the case of the Buck juvenile observed by Kristin, the bird was moving from a small lake with limited food resources to a much larger lake with a more abundant prey base. So the short-term benefit of being able to capture more food and put on weight for migration might be a large part of the explanation. Catching and consuming new food items might pay benefits down the road for young loons, as well, as this forces them to practice new modes of foraging that might be useful during migration and winter, when they will be faced with vastly different prey.

LMG_2823 Three Month Old Chick

There is a third and more nebulous hypothesis that might explain wandering by juveniles in fall. Three to four month-old individuals might be laying the groundwork for their lives as adults by exposing themselves to many different lakes and learning which are most suitable for them. That is, juveniles might try out lakes of different sizes and shapes containing different fish and invertebrates as a means of finding out what sort of lake allows them to feed most efficiently. If so, this period of exploration might allow them to target lakes more effectively when they return to northern Wisconsin (as they do) to look for a breeding territory in 2 to 5 years. It is even possible that young loons — especially males, which often settle to breed within a few short miles of their natal lake — move from lake to lake in order to create an internal map of the local area, which they will use in a few years when they return in adult plumage and seek a breeding territory.

Like many behavioral hypotheses concerning free-living animals, these 4 hypotheses for fall wandering are vexing to test. But they are not wholly untestable. For example, if juveniles simply move for better foraging, those that move should have higher masses in the fall than those that remain on their natal lakes. And if shifting between lakes prepares juveniles for foraging during migration and winter, frequent-shifting juveniles should enjoy higher survival than infrequent shifters. Even the hypothesis that juveniles begin to map out the local area for their later use can be tested to a degree, as it predicts more rapid settlement by frequent lake-shifters than infrequent ones. So we may, in time, begin to understand juvenile shifting. (This will come as a great relief to Kristin, no doubt, as she is braving the cool, rainy weather to document lake shifts by juveniles that we banded in July and early August.)

As I have noted before, the more immediate reason for tracking local movements of young juveniles has to do with learning about the development of natal-site matching behavior that we found recently. (Natal-site matching refers to the striking tendency of young loons to settle as adults on lakes that are similar in size and water chemistry to their natal lake.) We are curious to see whether loons show preferences for natal-like lakes even in their juvenile movements. If so, short trips during the fall to natal-like lakes will, of course, reinforce natal-site matching and yield young adults that have very strong natal preferences indeed!

Thanks, as so many times before, to Linda Grenzer, who provided these nice shots of the 2014 chicks on her lake. Although it is bittersweet for her, Linda hopes that these two juvies will soon themselves fly off to nearby lakes to gain valuable experience — or maybe just weight — that will prepare them for migration and beyond.

 

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