Tuesday, March 29, 2016

Do bald eagles delay incubation?

2014, Egg #2. Temp with windchill: -40F.
Do bald eagles delay incubation? It wasn't an a question we've thought about much, since bald eagles in Iowa usually lay eggs in temperatures under - sometimes well under - freezing! However, this year was quite a bit warmer, and the eagles in Decorah and Fort St. Vrain seemed to spend more time off the first two eggs than we are used to.

The question seems pretty cut and dried. After all, peregrine falcons delay full incubation very consistently until the third egg in a four-egg clutch, and the fourth egg in a five-egg clutch. In cold years (2013, 2014), we tend to experience higher losses than in warm years, since eggs one and two can freeze and die prior to the beginning of full incubation. However, it isn't that simple with bald eagles and many other birds.

2016, Egg#2: Temp no windchill: between 25-40F.
According to Professor Jim Grier, birds often start incubation slowly or gradually when temperatures aren't in the freezing range. Freshly laid eggs can spend a lot of time in the zone of suspended development (roughly 28.4 to 80.6°F or -2°C to 27°C) with no harm to the egg or embryo. While I don't have storage times for bald eagle eggs, Wikipedia tells me that the freshly laid eggs of domestic fowl, ostrich, and several other species can be stored for about two weeks when maintained under 5 C. That is quite a bit longer than the time between eggs one and two in a bald eagle nest!

Temperature aside, why do some birds begin incubating immediately? In a blog on the subject, David Hancock discussed two theories of immediate incubation: one, to protect eggs from marauding ravens and other predators, and two, to assure the survival of at least some young during periods of food scarcity. In the last case, the 'strategy' assumes that it is better to have one or two surviving children than no surviving children at all.

However, there are more benefits to delaying incubation so eggs hatch closer together: indeed, the survival of many waterfowl depends on it since adults leave the nest 24-48 hours after hatch begins, whether all of the eggs are hatched or not. Perhaps most importantly, delaying incubation helps assure that rapidly growing young are in the same stages of development. In the presence of an abundant food supply, more young will likely survive when incubation is delayed, since there will be less variation in size and ability between older and younger siblings, and food should be shared more equitably as a result.

Professor Grier pointed out that the nature (if not essence!) of biology is variation. This caused me to think back on eagles and monogamy. When we first started this adventure, everyone 'knew' that eagles were monogamous and mated for life. A new mate would be taken only if the old mate died. Remember when that was true? But once we really started looking, eagle relationships turned out to be more complicated. While many eagle couples appear to be monogamous, we've also documented extra-pair copulations, eagle 'divorce', polygyny, and polyandry.

The same is undoubtedly true for incubation. Professor Grier wrote that he would expect variation among different pairs of bald (and other eagles) in how much time they spent off the egg(s) both at the start and as incubation proceeded, as well as under different conditions such as temperature, humidity, and precipitation. We've certainly seen that in Decorah. In 2014, February's mean temperature was 9.4F - the coldest in 20 years of recording! Under those conditions, Mom and Dad incubated almost constantly. But in 2012, February's mean temperature was 27.8F - not the warmest ever recorded, but quite close to it. As indicated by video footage and all of the blogs and posts we did on the subject, Mom and Dad spent much more time off the eggs in 2012 than they did in 2014. While weather is clearly a factor, David Hancock noted timing variability between rural and urban bald eagles, and speculated that the presence of predators might make a difference in whether incubation was delayed.

The world is not a static place. Animal populations fluctuate as food supply, weather, and disease cause cycles of booms and busts. High populations and/or low food supplies can result in massive dispersals or irruptions as birds compete for territory and food. Populations may adopt different behaviors at different levels of density and food abundance, perhaps becoming less aggressive in a situation where neighbors and food are both in high supply. Everything changes over time: given a large enough timescale, the continents themselves flow like water. In that light, it is no surprise that eagle incubatory behavior might vary, or that eagles might adopt different behaviors as their population and/or the world around them changes.

So philosophy aside, can we really answer the question of delayed incubation in bald eagles and other birds? That's a tough one. The onset of incubation doesn't seem to involve timing in most species of birds, since they appear to be responding more to external factors than to an internal clock. This makes it hard to define what exactly 'delay' is for most birds. If I'm staying off my eggs because it is warm out, am I really a delayed incubator? If I almost always incubate right away because I lay eggs in sub-zero temperatures, am I really an immediate incubator?

Short answers are often necessary in our instantaneous and highly-connected world, so here is mine: "Bald eagles may or may not begin incubation immediately after the first egg is laid, depending at least in part on local factors that include temperature, humidity, precipitation, and predators. Apart from that, bald eagles exhibit variability: that is, not all pairs act the same way, even given the same or similar situations. There is more to learn, so keep watching, keep documenting, and trust the eagles!"

I'm going to close by quoting Professor Grier again: "The nature (if not essence!) of biology is variation." In my experience, almost every time we say a species always does this, or never does that, the species proves us wrong! Onset of incubation and mating systems are just two more examples of that.

Our ideas about birds have been shaped by how we observe them. Professor Grier pointed out that in the past, normal incubation details for various birds were studied from blinds, long-term and time-lapse photography, and with electronically telemetered artificial eggs, nest thermometers, treadles or scales to measure adult presence on the eggs. He wrote: "Eagle cams provide a whole new opportunity to observe and study traits such as incubation behavior (by individual parent, gender, total time, and under various environmental conditions) on increasing sample sizes of birds under undisturbed conditions." In other words, nest cams give us an unparalleled opportunity to observe large populations of birds in the wild, relatively free from human interference. Who knows what we will find, or what theories and beliefs will be challenged? We've got the cameras in the field. The next step is to develop appropriate methodologies and tools to enable large scale data collection across species and populations.

Thanks to the following for helping me learn about this subject.
  • Personal communication, Professor James Grier. Any mistakes I made in interpreting his email are my own. 
  • Personal communication, Bob Anderson. Bob used to tell us that eagles were pretty smart, but peregrine falcons were only about as smart as garter snakes. Even though he loved falcons, he believed that their lower intelligence made them less variable in their response to external stimuli. While that was outside the scope of this blog, I will return it later on.
  • David Hancock's blog on Bald Eagle Laying and Hatching Sequences