Monday, August 19, 2013

Eating and Evolution: Are Prey Preferences Causing the Evolution of Killer Whales?

When I was an undergrad, a lowly freshman who just knew she wanted to study biology, I took an internship at SeaWorld Orlando. I was excited that I got to participate in a real research project doing actual sciency stuff. The project was on the nursing behaviors of captive baby killer whales. Really cool right? Little did I know that actual science is composed of hours upon hours of tedious observation and documentation (2:00pm – melon bumping, 3:00pm – melon bumping, 4:00pm – melon bumping…). Despite that (or, who knows, maybe because of it), it was an neat project that boosted my interest research biology. And I got to watch killer whales for hours every week. So when I came across the paper for today’s post it really reminded me those times.

A new study published in the Proceedings of the Royal Society B, Biological Sciences looks at niche variation within sympatric killer whale populations in the North Sea. Those of you familiar with the terminology I just used might want to skip to the next paragraph. Otherwise, let’s hit a few terms first. We’ll start with the niche variation hypothesis. In the simplest terms, a niche describes where a species lives and the roles it plays in its habitat. The niche variation hypothesis describes differences within a species that are correlated with the variety of foods and habitats that are used by various populations. For example, why do island birds of the same species have different bill sizes? Likely because their bills adapt to the food items they are exploiting on their own island. It conveys a competitive advantage which results in a reproductive advantage that will lead, eventually, to an evolutionary change. This change will likely be a speciation event. This is a lineation-splitting event that produces two or more separate species from one (think about the branching on the tree of life). Usually we think of speciation as occurring via a geographic isolation (birds on different islands, populations separated by a mountain range, etc.), but the niche variation hypothesis allows for sympatric speciation because the exploitation of different resources splits a population within the same habitat. Admittedly, this type of selection would need to be really strong and stable over a long period of time to cause speciation. Now on to the study!

Killer whales (Orcinus orca) are actually members of the dolphin family (Delphinidae). They are the most widely distributed cetacean species in the world and are top marine predators. Males typically live about 30 years on average, and females about 50 years. The diet of killer whales is often geographic or population specific. Populations of orcas are usually defined as either “residents” or “transients.” As the name suggests, residents tend to stay in a more localized area whereas transients travel over large distances, sometimes overlapping with the ranges of resident populations. It has been documented that these different types of populations vary greatly in their diets, each consuming a narrow range of prey. Residents feed primarily on fish while transients feed nearly exclusively on other marine mammals. Considering this, and what we know of how the niche variation hypothesis cause speciation, have or are killer whales branching in to two species?

One of the problems in answering this question is the long-lived nature of these animals. It’s difficult to see a long-range change on a long-lived species. Most evolutionary studies use either comparisons at a single point in time or over timescales representing one to a few generations. Okay, that’s pretty good, and these snapshots have been very informative, but to get a real-time view in a long-lived species you really need to go small. And by that I mean molecular. Ancient DNA (aDNA) and stable isotope data from subfossil (remains that have not completed the fossilization process) specimens can be used to track niche and evolutionary history. The scientists in this study used these methods to look at the evolution in sympatric killer whale populations in the North Sea. First, they sampled 23 subfossil killer whale bones and teeth recovered by dredging or trawling the Southern Bight of the North Sea or from archaeological sites in Southern Scandinavia. Then they dated their samples using radiocarbon techniques or archaeological context. Next, they used stable isotope ratios to provide a long-term measure of what the animals ate during their lifetimes and thereby estimate the orcas’ niche width (it is argued that populations in wider niches are more variable than populations in narrower niches). Additional evidence of these dietary habits was gathered from examining the wear-patterns on the teeth (for example, feeding heavily on herring badly wears down the teeth). Then mitochondrial DNA (mtDNA) sequencing was used to determine the degree of linage sorting (separate populations carry their genetic diversity with them) based on isotopic (prey) niche. And finally, they biopsied the skin of modern orcas, sampling either while the animals fed on fish or on stranded remains with known stomach contents. From this they were able to extract high-quality DNA and conduct an individual-based analysis of population structure. This, combined with the aDNA data, effectively gave them a map of the evolutionary outcome of niche variation.

This is one of those studies where the results are all variable. *sigh* ‘Tis science. From the isotopic analysis, the researchers found  a lot of overlap in the results, mostly likely explained by among-individual differences. Because this type of analysis represents what an animal ate over its lifetime, differences in prey items within the diets of individuals are not apparent. This and the analysis of tooth wear suggests some overlap either in the diet and/or foraging method of the specimens studied. The result is consistent with the observations of the modern whales. Fish eating pods are often found with mammal remains in their stomach contents. Lineage sorting of mtDNA sequences based on the isotopic values revealed that there “has been multiple diversifications [sic] in isotopic niche” and “an indication there was relatively stable transmission of isotopic niche along matrilineal lines within some clades, in particular those that were dominated by samples from Norway.” The incomplete lineage sorting they found seems to be consistent with relatively recent divergences in niches, and their models indicate panmixia (random mating) between at least some groups that feed on fish and some groups whose diet includes seals.

To sum up, we know that there is niche variation in populations of killer whales. But all of that variation and overlap that the researchers found suggests that any speciation is still at an early stage in this system. And while the results of this study seem to be all over the place, it does add more information to the story while providing a useful long-term evolution study methodology. It also strengths the argument that sympatric speciation is difficult to achieve.

Also check out this great presentation on this study!

ResearchBlogging.orgFoote, Andrew D., Newton, Jason Newton, Ávila-Arcos, María C., Kampmann, Marie-Louise, Samaniego, Jose A., Post, Klaas, Rosing-Asvid, Aqqalu, Sinding, Mikkel-Holger S., & Gilbert, M. Thomas P. (2013). Tracking niche variation over millennial timescales in sympatric killer whale lineages Proceedings of the Royal Society B, Biological Sciences, 280 (1768) DOI: 10.1098/rspb.2013.1481

Science's article "North Atlantic Killer Whales May Be Branching Into Two Species"

For more information and explanation of some of the evolutionary terms discussed this post see:
Understanding Evolution via Berkeley, particularly the page on sympatric speciation
and for a nice description and examples of niche variation see
Soule, M. and Stewart, B.A. (1970) The "Niche-Variation" Hypothesis: A Test and Alternatives. The American Naturalist, 104(935): 85-97. (LINK)

Some useful resources for information on killer whales:
NOAA Fisheries Office of Protected Resources page on Killer Whales
National Marine Mammal Laboratory's page on Killer Whales
Cascadia Research Collective's "Studying the diet of fish-eating killer whales"

(image via National Geographic, photo credit Gerard Lacz/Animals Animals—Earth Scenes)
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