I began studying KIMU breeding biology in Kodiak, Alaska, in 2008 as part of a conservation effort to determine the causes of a dramatic population-wide decline that had been detected by boat-based surveys. Through the course of my research I have used the extreme phenotypic differences and the evolutionary relationships apparent in the Alcidae, as well as inferences made from patterns in convergently evolved species, to make predictions about emerging aspects of the breeding ecology of KIMU, and make hypotheses about this intriguing species.
Kittlitz’s murrelet has been the enigma nonpareil among the alcids, and a large part of this owes to the extreme difficulty researchers have had in finding their nests. They breed non-colonially in rugged alpine areas, nest up to 75 km inland (where predators are often abundant), are cryptically colored, and they are extremely secretive near their nest sites (Day, Kuletz, & Nigro, 1999). Its habits are so mysterious and its Beringian alpine breeding habitats so remote that only 25 nests had been described prior to 2005. I used the smattering of anecdotal data from these previous nests, along with information about its relatively well-described kin, the marbled murrelet (MAMU), to design an expedition that would yield a large number of nests. My team knew about the close evolutionary relationship between the KIMU and MAMU from phylogenetic relationships determined from DNA and morphological analysis (Cracraft et al., 2004; Friesen, Baker, & Piatt, 1996; Moum, Johansen, Erikstad, & Piatt, 1994; Strauch Jr, 1985) (Figure 1, Friesen et. al, 1996). Since the two species appeared to have diverged only about 2 mya, I inferred that the breeding biology of the two species would be much more similar than a comparison between KIMU and other clades of the Alcidae, since many of these clades radiated from a common ancestor about 18 mya, and would be expected to have independently evolved many derived traits that would not be shared with the KIMU. Using MAMU as a proxy species, I made the predictions that: 1) KIMUs nest in a very dispersed manner on the landscape to avoid predation; 2) KIMUs exhibit nest area fidelity, meaning that pairs usually return to the same place every year to breed. I also made predictions based on the commonalities among the dispersion patterns of other cryptic species such as grassland birds and peppered moths to infer that these groups convergently evolved their dispersion strategies (i.e. these species are well-spaced throughout their potential habitat) to avoid predation.
So, following these predictions, I prepared an expedition that could move great distances and cover as large an area as possible to account for high dispersion of nests, and that would be reproducible in following years to find birds that were nesting in the same area annually. The effort turned out to be a success. By spending up to three months at a time backpacking among likely alpine habitats, my group discovered 53 active KIMU nests during 2008-2011, generating a wealth of new information about the breeding ecology of the species. And, as is usually the case in science, we also came up with a lot of new questions.
The first and most obvious question we had is: why are these birds doing such a poor job of reproducing? Only 9 of 53 (17%) nests produced a fledged chick, a very low figure that has recently been corroborated by other studies in different areas of Alaska. Again, looking at phylogenetic relationships we saw that most of the Alcidae experiences much higher rates of nest success. We have no idea whether our small window of data on KIMU is representative, but we do know that the rarity of the species, and the relatively small area of breeding habitat available to it, that it may never have been abundant in recent history. “Recent history” is the operative phrase. The combination of KIMUs affinity for glaciated habitats, and for foraging in glacial outflow areas, and it’s apparent emergence from a common ancestor with the MAMU 2 million years ago, suggests that KIMU may in fact be an ice age relic species, adapted to a eking out its nesting period on bare mountaintops surrounded by glacial ice, as well as in the scree and talus fields created by retreating glaciers. Such habitats would be much less common today than they were during the major glacial retreats that occurred about 10,000 years ago, when large areas of Alaska were melting out from a blanket of ice several kilometers thick. Certainly this may only be circumstantial evidence, but it’s tempting to think that the shrinking glaciers of today may be leading to the shrinking population of KIMUs as well. This hypothesis could be tested, though at great expense, from a large scale study of nest sites and their relationships with glacier dynamics.
But how long do KIMUs live? If their life spans are long enough, even a 17% nest success rate would be sufficient to sustain the population. As you might expect, however, we have no clue about the average adult mortality of KIMU, so we go back again to the phylogenetic drawing board to determine how long its closest relatives typically live. The results are a little sobering: a 15 to 20 year average life span is what is predicted for MAMU, a species that also happens to be in decline through most of its range. Though models of MAMU survivorship and nest success can’t be expected to fully explain KIMU population dynamics, they do provide our best guess as to how long KIMU adults would need to live on average for population stability, and our best models suggest that 15-20 years may not be long enough.
It’s also tempting to compare the breeding strategy and plumage of the KIMU with its alcid kin. Since most of the Alcidae are colonial, nest very close to the sea, and exhibit countershaded coloration, we could presume that these traits are somehow adaptive. KIMU does not share these traits, presumably because of trade-offs resulting from nesting in predator dense areas: it would be a liability to have contrasting coloration and dense colonies where foxes, for example, may be abundant. As a result of these trade-offs, KIMUs may need to spend more time and energy foraging and provisioning fish than other alcid species, meaning that changes in food supply, water temperature, and other factors may have a disproportionate impact on KIMU, relative to other alcids. Again, this is a testable hypothesis, one that I am exploring by studying the diet composition and growth rates of KIMU chicks.
The patterns of evolution have been fundamental for making predictions and hypothesis for a seabird that whose breeding habits were formerly almost completely unknown. Not only have I used phylogenetic relationships to make informed hypotheses relating to dispersion and breeding ecology of KIMUs, but managers will use inferences informed by phylogeny to make educated guesses about life history parameters such as adult mortality for which data does not yet exist. Far more than a merely academic pursuit, principles of evolution may help us to ensure that this cagey little seabird will continue to be found for a long time in the Alaskan alpine gnar. If you look hard enough, that is.
Cracraft, J., Barker, F. K., Braun, M., Harshman, J., Dyke, G. J., Feinstein, J., Stanley, S., et al. (2004). Phylogenetic relationships among modern birds (Neornithes). Assembling the tree of life, 468–489.
Day, R. H., Kuletz, D. J., & Nigro, D. A. (1999). Kittlitz’s Murrelet (Brachyramphus brevirostris). In A. Poole & F. Gill (Eds.), The Birds of North America. Philadelphia, Pennsylvania: The Birds of North America, Inc.
Friesen, V., Baker, A., & Piatt, J. (1996). Phylogenetic relationships within the Alcidae (Charadriiformes: Aves) inferred from total molecular evidence. Molecular Biology and Evolution, 13(2), 359.
Moum, T., Johansen, S., Erikstad, K. E., & Piatt, J. F. (1994). Phylogeny and evolution of the auks (subfamily Alcinae) based on mitochondrial DNA sequences. Proceedings of the National Academy of Sciences, 91(17), 7912.
Strauch Jr, J. (1985). The phylogeny of the Alcidae. The Auk, 520–539.
I think it is fascinating how biologists and other scientists can take information as varied as ice age topography/geology, phylogenetic data, and historical accounts, and use it all to make predictions about a rare bird (or even where you're likely to find fossils--i.e., Tiktalik). Granted, science is all about predictions and investigating how those hold up to various scenarios, but there's a difference between predicting that a ball dropped from a certain height will hit the ground at a certain velocity and such "abstract" areas as determining the breeding habits of a rare bird (though I guess if that's your area of research, it's not very abstract anymore!).
ReplyDeleteYou mention that the KIMU might be a "relic" of the ice age, because of its natural habitat and how it seems adapted to live there. Due to the increasing shrinkage of their habitat, you also imply that the species may become extinct if circumstances continue as they currently are (given the estimate of adult life span, per the MAMU data, is rather low as compared to nest success rate). How feasible do you think it is for conservation efforts to sustain the KIMU population, despite environmental/human factors?
It is quite impressive what biologists will be willing to do in order to get a better look at their subject of interest. I think someday I would like to be a part of an expedition like the one you planned out and went on in order to collect data on a certain species of animal. The thought of backpacking in a glacial setting of Alaska sounds frightening and exciting at the same time, but I can tell yielded your study valuable information that you could not have collected any other way. I was wondering, how much planning goes into creating an expedition like the one you went on?
ReplyDeleteI also think it is amazing that you can use all the concepts of phylogenetics in order to make reliable predictions about an otherwise not well-known species. The fact that you deduced that KIMU was an “ice age relic species” based off of its divergence from its common ancestor 2 mya where glacial outflows and glaciated habitats were more common is a very well-thought out conclusion. A question I have in regards to this is, what do you think is going to happen in the next ten years or so to the KIMU population if the glacial ice continues to recede? Will they learn to adapt in some way? Or will their dependence on a certain habitat and propensity of a low fledging rate get the best of them?
This is wonderful work; congratulations on your success to this point in finding so many active nests for such a rare species. From just reading this I agree that this species seems to have tucked itself into this very tiny niche (nesting in close proximity to glacial outflow). Considering the wide phylogenetic and behavioral gulfs between the KIMU and the other alcids there is obviously some reason the KIMU have not made the adaptations their closest relatives have. The stress inflicted by nesting so far from the ocean would seem to be so great that one has to wonder why the KIMU started down this path in the first place; it has got to be part of the answer as to why the KIMU are having such poor breeding success. Do any other researchers have information (particularly fledge rates) for birds that are nesting near glaciers that are relatively close to the ocean?
ReplyDeleteUnfortunately, the writing seems to be on the wall for these cool birds. Glaciers are not going to be advancing anywhere, any time soon, and the situation is probably going to get much worse over the next few hundred years. The fact that these birds are obviously straddling two completely different environments (webbed feet and flying underwater are a remarkable contrast to nesting on scree slopes near glaciers)does not, I think, bode well for the possibility of their changing their lifestyle to the point where they will survive long-term. I hope the work on them will continue and that we will pay attention to whatever the KIMU can tell us.
This study sounds fascinating, and the blog was very well written. I loved the introduction; it was both informative and humorous! The expedition sounds amazing, like a biologist's dream. I think it is really great that although it is challenging for researchers to study and really understand this species because of the difficulty in finding nests, you and your team used past data and were able to track as many nests as possible. How long did it take to acquire the necessary information to successfully find the nests? What were the difficulties and drawbacks of planning an extensive expedition like this?
ReplyDeleteI think what particularly drew me to this study was how environmental factors like the shrinking glaciers may be affecting the reproductive and survival success of this species. It is a sad state of affairs that not only are melting ice caps and glaciers affect the species that directly rely on them for refuge, but that the melting of these ice caps are also affecting our oceans, which causes sea level rise, ocean temperature changes, declines in marine species, and what has recently been studied, changes in ocean currents. The consequences are innumerable. Unfortunately, it seems like due to their decline in population and their common ancestry to the MAMU who live in glacial ice, the KIMU are trying to survive in an area that has change drastically from the environment they are supposed to survive in. It is good to know that even with the small nest success rate, the populations are able to survive because of possibly a long lifespan. What are long-term plans for the KIMU? Do scientists know what actions will be taken if the populations decline so low that this species cannot successfully survive in the wild anymore?
I really enjoyed reading this post. I have never heard of this bird before, but I have worked with various seabirds in the past and find them to be very smart and quite interesting. Your research on KIMUs must be 'so exciting, especially going on the expedition! It is amazing that your prediction on KIMU nesting sites based on their closest relatives and other cryptic nesting birds was so successful and that you added so much knowledge to what is currently known about the species. I would like to know more detail about how you were able to use that information to locate very specific spots and from there how you were able to study the birds.
ReplyDeleteOn the topic of glaciers, from your research did you find any evidence of the KIMU as evolving to meet the fast changing environment?
Also, what kind of impact does the KIMU have on its environment? Other people have asked what will happen to the species if it declines so low that it would be basically impossible to restore the species. If the KIMU became extinct what impacts would that have on the environment?
It’s amazing what you can get from some anecdotal data and hard work connecting the dots. Congratulations on the successful expedition which while bringing up many more questions will at least give you a place to start as you begin to answer some of these questions. You mentioned that because of trade-offs that KIMU is more sensitive to changes in food supply, water temperature, and other factors. How do you see global warming impacting KIMU in the short term and long term? It seems this cagey little seabird is definitely a fighter but with it already in a fragile state with only 17% of nests producing hatchlings it begs the question, will species like this survive?
ReplyDeleteKIMU seems to be a species that like someone else mentioned is a “relic of the ice age”, how do you feel about that? Is its fragile state simply a consequence of the times that it’s in? Do you see KIMU making a comeback should we face an ice age or do you foresee greater issues affecting this species? What has your data from the study of diets told you about the effect of global warming on the diet of these birds? What about environmental pollution?
I wish you the best of luck with your research, it is definitely an interesting organism to study and I hope to hear more in the future.
Anatoliy Vlasenko