[I tried sending a version of this note earlier today, but it didn't go 
through; I apologize if the earlier version re-emerges from the ether.]

The question is not whether any hybridization at all is occurring, but whether 
hybridization is a plausible explanation for a major change in the average 
appearance of an entire taxon.

>To me it makes no more sense to dismiss hybridization amongst those two 
>species as a mechanism for primary darkening than it does to invoke it.

The hypothesis should not be dismissed, but nor should it be invoked 
uncritically. Instead, let’s do our best to evaluate it on its merits. Despite 
all the deficits in our knowledge, I believe that the hybridization hypothesis 
can be challenged on two fronts: (a) the mechanism’s theoretical ability to 
produce the effect in question (i.e., a major increase in primary pigmentation 
across an entire taxon); and (b) the extent to which we are observing the 
corollary consequences that would be expected if this mechanism were operating 
at a scale sufficient for its efficacy in (a).

Regarding (a), although gene flow is one of four mechanisms that can alter gene 
frequencies in a population (the other three are mutation, selection, and 
random drift), it is not by itself expected to cause huge changes--for instance 
to cause a trait that was previously common to become rare. Much more 
typically, it might introduce a new allele into a population (i.e., raise this 
allele’s frequency from near zero to slightly above zero), thereby potentially 
allowing natural selection to gain traction if the allele proved favorable in 
the new population. Gene flow could easily add some dark-winged birds to a 
white-winged population, thereby causing the white-winged trait to drop from, 
say, 100% to 99%; but in the absence of selection, astonishingly high rates of 
hybridization would be required to transform a whole taxon from >90% 
white-winged to >90% gray-winged—rates that are not only improbable in 
themselves, but which would also produce many other observable consequences.

> A Western birder might find the idea that two similar gull species *not* 
> hybridizing to be alien to their experience, for example.

Consider any example of hybridization that you like. One thing that you will 
notice is that wherever the hybrids are at all common, at least one (and 
usually both) parental types will also be common. Wherever one finds lots of 
Mallard x Black Duck hybrids, one finds hordes of Mallards and Black Ducks. 
Even at the epicenter of Glaucous-winged x Western Gull hybridization, 
Glaucous-winged Gulls are abundant in migration. This brings us to point (b). 
If hybridization with Thayer’s Gulls were occurring on the massive scale 
necessary for flipping a trait value in North American Iceland Gulls, one would 
expect to see a lot of Thayer’s Gulls wherever the hybrids (i.e., Kumlien’s 
Gulls, under this view) were common. This is manifestly not the case along the 
New England and Long Island coast, where the frequency of convincingly 
Thayer’s-like birds is so close to zero that it is actually shocking—one would 
think we’d see more than we do strictly via vagrancy, even if there were zero 
hybridization going on.

Heavily pigmented Iceland Gulls are not in any way anomalous. They are the norm 
in coastal northeastern North America, and their increasing occurrence in 
Europe may be due to vagrancy, or it may be due in part to gene flow and 
natural selection acting on Greenland-breeding populations. What reason is 
there to doubt that, by and large, these pigmented birds return north and breed 
with each other, that their offspring are also pigmented, and that they all 
continue to live and act (and sound—thanks, Pete) like Iceland Gulls? The 
alternative is to posit that, upon returning north, the palest ones are 
systematically knocked up by randy, vagrating Thayer’s Gulls that, despite 
their depraved penchant for miscegenation, still assiduously conform to 
traditional Thayer’s Gull migratory routes, eschewing the wintering areas of 
their bewildered mates and progeny!

Shai Mitra
Bay Shore



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