A few days ago, I was conducting observations on the lake where Linda Grenzer lives when she snapped this photo of me, the two pair members on the lake (the two loons closest to my canoe), and five territorial intruders. Linda’s picture captures beautifully what transpired during my visit of an hour and a half. Better yet, the photo shines a spotlight on an enduring question that drives much of my research on loon territoriality: what is the purpose of territorial intrusion. This is a question that we have only half-answered.
We do know that many intruders visit in order to learn whether or not a territory has chicks; if they detect chicks, they are more likely to intrude in the following year and attempt to evict the breeding pair member of their sex. (This makes some sense, as the presence of chicks is an indication of good nesting sites and plentiful food for young.) We also know, from a recent analysis, that loons practice natal-site matching; that is, they attempt to settle as a breeder on a lake that is similar in physical size and in pH to the lake they were reared on. So undoubtedly some intruders must be learning about lakes where they intrude so that they can settle on one similar to their natal lake.
But there must be more motivation for intruders to visit lakes defended by breeding pairs. Among the 5 intruders pictured in the photo, for example, were: a banded 9 year-old loon not known to be settled on a territory yet (i.e. a probable “floater”), an unbanded loon whose status is wholly unknown, and three banded loons known to be members of breeding pairs from neighboring territories. While the first two birds could plausibly be shopping for territories through chick detection or natal-site matching, the neighbors are unlikely to be doing so.
What, in fact, could neighbors stand to gain from intruding next door? Several hypotheses are possible here. Neighbors might gain by becoming familiar with other loons with which they might become mated in the future, if they both lose their current breeding positions and must settle elsewhere in the general area. Neighbors might also be trying to learn about the territory, which they might occupy in later years, providing one of the current pair members dies. Another possibility is that neighbors have no particular interest in the territory where they intrude but, rather, are intruding in order to draw attention away from their own territory (since loons on the water tend to attract flying loons to land and investigate). That is, intruders might be attempting to “decoy” loons away from their own territory so that others do not learn about it and attempt to settle there. Finally, neighbors might simply visit to forage in someone else’s territory, depleting the food supply there instead of at home.
An additional question raised by the photo is: what is the breeding pair’s response to intruders? One might expect that the breeders would react aggressively towards intruders, driving them out immediately so that they cannot learn about their chicks (of which there were two in this case, hiding our near shore and far from the intruders) or harm them. Yet this pair — typical of breeders — was tolerant of the intruders and allowed them to roam freely throughout the territory for over an hour. Are the breeders feigning nonchalance to reduce the likelihood that intruders will look closely for and detect the chicks? Or do large numbers of intruders pose a severe threat to territory ownership such that territory owners must tolerate them or risk losing their positions?
I hate to raise so many questions that we cannot answer immediately. Testing of most of these hypotheses for intrusion and defense towards intrusion is feasible in our population. For example, we can look statistically at the “decoy” hypothesis by seeing whether pairs that vacate their territories and intrude next door experience a lower rate of territorial eviction than pairs that remain on their territories faithfully throughout the season. And we can test whether pairs that attack and stalk intruders, rather than tolerating their intrusions peacefully, suffer a higher rate of territorial eviction, because they betray the presence of their chicks and place a target on their own backs.
Such statistical analysis requires large samples of lakes and intrusions, so it will take time. Meanwhile, we will have to enjoy the experience of tracking intruders and breeders around territories by canoe and wondering what peculiar combination of evolutionary interests of breeders and intruders could produce such flotillas.