Juncos: What do we know?

From Mark Szantyr:

While I enjoy debating the geographic origin of first year Herring Gulls as much as the next guy, and hey, you can never see enough images of an American Robin in the UK (?), I wanted to change modes here for a bit and ask your opinions on a few juncos that I have photographed and banded. It seems that I am receiving numerous reports or various forms of "Oregon" junco this year and after inspection, except for one bona fide female Oregon-type, I have relegated all to a hyemalis origin even though they show some striking characteristcs not normally considered in the realm of pure J. h. hyemalis possibilities. My criteria has been largely rooted in the Alden Miller works. Am I being too limited in what I think an Oregon Junco, or Pink-sided Junco, looks like? So, I have posted images of two different birds ( I have many more to work through) on the Surfbirds North American Stop Press page. The address for Surfbirds is : http://www.surfbirds.com and follow the links to the North American Stop Press page. These are two birds from my studies over the past years of this intriguing complex. I have evidence to suggest I know the ssp of one of these birds. I believe I know the other as well but surly would like to entertain your thoughts. Please e-mail your suggestions on this topic to me directly or if the subject warrants we can discuss this in an open forum. What do we really know about the field ID of ssp's of Junco hyemalis?

Mark S. Szantyr, Connecticut (10 Jan 2004)

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From Matthew Kenne:

I'll take a stab at them to hopefully spur on some discussion. First the February bird: There doesn't appear to be any "western" hood on this bird. I'll stick with an immature hyemalis bird at the very brownish end of the spectrum. The October bird shows the hood strongly even though the gray is heavily mixed with brown. I believe the brownish overlay of the hood in younger or female birds is more a feature of the montanus (by Alden) or shufeldti (by Pyle) subspecies than of those farther west.

Matthew Kenne, Iowa (10 Jan 2004)

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From Ted Floyd:

Hello, birders.

Mark S. Szantyr writes:

> It seems that I am receiving numerous reports or various forms of "Oregon" junco this year and after inspection, except for one bona fide female Oregon-type, I have relegated all to a hyemalisorigin even though they show some striking characteristcs not normally considered in the realm of pure J. h. hyemalis possibilities.

The conventional wisdom, it seems, is that most eastern juncos are Slate-colored (hyemalis), with a handful of Oregon (oreganus) mixed in. My guess is that this belief traces back to the treatment of the juncos in the fourth edition (1980) of the Peterson Field Guide. It's *still* (by far) the most frequently used field guide, after all--and its various inaccuracies are slow to be extinguished.

An interesting exercise is to go back through eastern U.S. CBC data from the 1970s and 1980s and count up all the records of Oregon Junco. The bird is ubiquitous! ...

... 'cept it ain't. Banding data (e.g., from the Powdermill banding station in western Pennsylvania) confirm that Oregon Junco is exceedingly rare. Another possibility is Pink-sided (mearnsi); the complication here is that Pink-sided has often been treated as a subspecies within the Oregon subspecies-group--with the result that few birders have paid it much attention. (In the old days, for example, the CBC accepted records of Oregon, but not of Pink-sided). But we still have a problem: Good records of Pink-sided Junco (e.g., banded birds, specimens, or diagnostic photos) are extremely rare in the East.

So far, then, we have the following:

So, what are these birds? Well, presumably, White-winged (aikeni) and Gray-headed Junco (caniceps) aren't in the running.

They are, I believe, Cassiar Juncos. Now there's a name that I suspect not all of us would recognize! But it's for real. The Cassiar Junco (cismontanus) is regular in winter in the eastern U.S. At the Powdermill banding station, for example, some 10-15% of the juncos (about 35,000 juncos in total) have been Cassiar.

Cassiar Junco is treated extensively in Alden Miller's 1941 monograph, and it is depicted (without being named) on the bottom right of p. 501 of the Sibley Guide.

Compare the pictures on the Surfbirds website with the photos on p. 135 of the April 2003 issue of Birding. The accompanying article ("The junco challenge: A genuine Pink-sided Junco from Arkansas and some look-alikes", by Joseph C. Neal) is definitely worth the read.

Mark's Bird #1 looks especially good for Cassiar Junco. Bird #2 is a mess (what fun!), but I don't think I'd call it an Oregon or Pink-sided. It's got a fair bit of Slate-colored and/or Cassiar genes in it, I believe; and maybe all of its genes are Slate-colored and/or Cassiar.I wouldn't say that the bird definitely lies outside the range of variation for Slate-colored/Cassiar.

Something else: I may have harped on this point before, but the AOU really needs to confront the Cassiar Junco conundrum. The bird is currently classified within the Slate-colored subspecies group, I believe. But that doesn't make any sense. Cassiar Junco clearly is *intermediate* between Slate-colored and Oregon Juncos; depending on your views of evolution and speciation, it is either an "intergrade" or a hybrid, or even a "stable hybrid". But it is no more a Slate-colored Junco than a first-generation Brewster's Warbler is a Golden-winged Warbler. As long as we're going to recognize Cassiar Junco (which I think we should continue to do) and give it its own trinomial, then (a) we have a problem with calling it Slate-colored and, (b) we have a problem with distinguishing between Slate-colored and Oregon as separate subspecies-groups.

Finally, if I may, another remark about Peterson's treatment: If you actually read the text, Peterson basically get its right when he says that "intergrades between oreganus and typical hyemalis are frequent"; but then, frustratingly, he advises that we "just call them Northern Junco, now the proper name for the species". I can't imagine anything more dissatisfying than leaving all juncos unidentified! The more I look at and think about juncos, the more I realize that there are serious problems in our "classical" methods of giving names to populations of birds.

Best, Ted Floyd, Colorado (13 Jan 2004)

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From Alvaro Jaramillo:

Ted Floyd wrote:

Something else: I may have harped on this point before, but the AOU really needs to confront the Cassiar Junco conundrum. The bird is currently classified within the Slate-colored subspecies group, I believe. But that doesn't make any sense. Cassiar Junco clearly is *intermediate* between Slate-colored and Oregon Juncos; depending on your views of evolution and speciation, it is either an "intergrade" or a hybrid, or even a "stable hybrid". But it is no more a Slate-colored Junco than a first-generation Brewster's Warbler is a Golden-winged Warbler. As long as we're going to recognize Cassiar Junco (which I think we should continue to do) and give it its own trinomial, then (a) we have a problem with calling it Slate- colored and, (b) we have a problem with distinguishing between Slate-colored and Oregon as separate subspecies-groups.

Ted brings up an interesting point, mainly that folks are not necessarily aware of the existence of this beast, cistmontanus, the Cassiar Junco. If you head up to Jasper National Park in Alberta you see these things all over the place, as well as other birds that look like pretty good members of both Oregon and Slate-colored Junco. In Jasper you can also see hybrid Audubon's x Myrtle Warblers, and just south of there you can see the gambell's White-crowned Sparrow turn into Mountain White-crowned Sparrows. Clearly it is an area where several taxa are in secondary contact, and I am quite sure that this is what the Cassiar Junco is a product of. Not only is the distribution of the beast important in understanding how to assess the taxonomy of the Cassiar Junco, but also in the breeding ground its variability in plumage characters suggest it is of hybrid origin. So if this is what explains the Cassiar Junco, mainly that it is a hybrid population, then we cannot recognize it as a subspecies. It goes against the general rules of how to apply the subspecies concept. So I would suggest that we continue to use the English name "Cassiar Junco" to refer to these birds, but that we drop the name cistmontanus for this population. I don't know if Jim Rising is out there listening in on this, but it would be interesting to see if he agrees with this or not. I recall having many, many, many chats about Juncos with Jim when I was his student. This group is maddening, the organization of the different entities is a mess, and lumping them all as one was probably not the right thing to do, but there are so many options in how to divide up the group and differences in level of hybridization between the different forms that I understand why no one has decided to go ahead and figure this all out. As a related aside I think that the Fox Sparrow taxon altivagans could be a parallel situation with the Cassiar Junco, but it is less clear. It may be a hybrid population between "Red Fox Sparrows" and "Slate-colored Fox Sparrows".

Cheers, Alvaro Jaramillo, California (13 Jan 2004)

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From Ned Brinkley:

Juncos similar to the bird(s) that Mark Szantyr has posted on the Surfbirds website are known from Virginia, at least in recent times, and I've seen a few while working the junco flocks, but very few in the case of the second image posted (Feb 02). The first and third images depict a plumage that's seen relatively often; I have not assumed that these birds are anything other than young Slate-colored of some sort.

The status of Cassiar Junco in the state is unclear, but they would seem not to be as numerous as Ted Floyd notes for Pennsylvania, for instance. I've not had a chance to go over the state specimens carefully, and I'm not sure how well the older accounts' identifications of individual juncos (such as Murray 1952) dovetail with Alden Miller's descriptions. However, there are a few _cismontanus_ specimens out there from Virginia (Buckley, P. A., and F. G. Buckley. 1967. The Current Status of Certain Birds in the Virginia Capes Area

II. April 1967--July 1968. Raven 39: 27-40; a good discussion of juncos there) from southeastern Virginia. I have suspected that this is the form I've seen several times, most convincingly in Suffolk near Wilroy in early January 1997 (a bird that looked like Szantyr's central image, the Feb 02 bird). I will post photographs of specimens at some point -- the state sorely needs a re-evaluation of all records of potential non-Slate-colored sorts. (Apologies, by the way, to readers of North American Birds and to the authors of the recent piece on vagrants to Long Island -- the images of the junco reproduced rather poorly in the journal, being quite washed out -- not very helpful when dealing with subtle distinctions. The inexpensive paper used for the journal is unpredictable at best.)

Ned Brinkley, Virginia (13 Jan 2004)

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From Wayne Weber:

Bird ID People,

I believe that in many ways, Ted Floyd has hit the nail on the head in his discussion of the "Cassiar Junco". This form (considered a subspecies of Slate-coloured Junco in the 1957 AOU Check-list) breeds over a wide area of northern British Columbia and the southern Yukon, and in a relatively small area of mountainous western Alberta. (Cassiar, by the way, is a now-defunct asbestos mining town in the far northwest of British Columbia.) However, Ted and Alvaro are probably correct that the "Cassiar Junco" is really an intergrade or hybrid population, rather than a subspecies in the usual sense.

Because of their intermediate characters, I suspect that Cassiar Juncos tend to be labelled as "Oregon Juncos" when they show up in winter in the East, and "Slate-coloured Juncos" when they show up west of the Rockies in southern B.C. or southward. They look different from the typical Slate-coloureds in the East or Oregons in the West, but don't have all of the features of the other form.

In fact, I suspect that the existence of the "Cassiar Junco" is alarge part of the reason why the Slate-coloured and Oregon Juncos were "lumped" in the first place-- although the lumping didn't stop there,but went on to include all the other North American juncos except theYellow-eyed.

Ted is also correct, in my opinion, that "the AOU really needs to confront the Cassiar Junco conundrum". Not only that, but the Committee on Classification and Nomenclature really needs to confront the entire question of subspecies, which it has not done since the 5th edition of the Check-list in 1957. We now have an up-to-date treatment at the species level of bird classification for all of North and Middle America (the 7th edition, published in 1998), but we are stuck with a 50-year-old treatment of bird subspecies. Without belittling the magnitude of the task, current conservation issues, as well as scientific ones, make it imperative that a modern treatment of subspecies be completed soon.

However-- whatever treatment for juncos the AOU Committee eventually decides on, I suspect that the Cassiar Junco will be a thorn in their sides. Evolutionary patterns are not always neat or easily explained!

Wayne C. Weber, British Columbia (13 Jan 2004)

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From Ian McLaren:

All:

It is good to see the current thread on juncos, something I've been pondering for some time. In Nova Scotia we have 20 records of "Oregon Junco" in print, with almost none verifiable from the terse descriptions. One was collected years ago: I'll look into it some time.

It is clear, however, that of a half dozen photographic records examined by me, only one is a certifiable "Oregon." That one is was a singing male (!) cf. _montanus_ photographed 10 May 1997 on Seal Island. One is arguably _mearnsii_ (differing expert opinions), and the rest are "Cassiars."

I'm glad to see the suggested retirement of _cismontanus_. Dave Sibley captured the views made here by others by designating the Canadian Rocky Mountains birds as a "broad intergrade population." (As others have noted, arbitrarily assignment of _cismontanus_ to the "Slate-colored group makes little sense.) He might have unintentionally added some confusion, however, by showing a very similar brown adult female under Slate-colored. That bird, too, would seem well within any "broad intergrade population."

I have made it a point to look carefully at early fall juncos here, and have never seen a bird as extreme as this "brown adult." I believe that the late autumn or winter appearance of such birds here - some have been "ticked" as Oregons as noted above - is an indication that they arrive from afar. That is, I don't believe birds as brown as Dave's adult female are produced in these parts. What do others know of the timing of appearance of such birds in, say, the mid-west?

Another point is the designation of some eastern records as _J. h. (o.) oreganus_ in some works (e.g. Birds of Massachusetts), although that Paicific slope race probably rarely if occurs in the East. "Good" Oregon-group birds in the East are presumuably _shufeldti_/_montanus_.

Finally, note that in one respect Mark Stantyr's bird in the hand is on the "Oregon" side, with a good, convex lower margin of its distinct hood, whereas the bird on the ground is messier in that respect.

Cheers, Ian A. McLaren, Nova Scotia (14 Jan 2004)

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From Kent Nickell:

We had a junco at our feeders in central Iowa this time last year that I thought might be an oregon but am wondering if it would fit into the Cassiar. I have it at http://www.greenbackedheron.com/photo.cfm?photoid=1393.

I also have an apparent April Gray-headed from Colorado that shows some white on the greater coverts at http://www.greenbackedheron.com/photo.cfm?photoid=1392.

Kent Nickell, Iowa (14 Jan 2004)

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From Mark Szantyr:

Thank you all for the responses so far...this has been really interesting and I hope to continue this discussion. I have a few more images I will load in the next day or so...let's call it Round 2 of Guess the Junco.

Why would Kent's image not be an Oregon Junco?

The previous two birds I posted on Surfbirds are rather interesting. The Feb 02 bird arrived at my feeder in late December and stayed through late April. Arriving as a very red bird ( as pictured) , the bird changed to arelatively normal pale gray hyemalis female-type by the the time it left (it was banded so I was able to follow its progress). The bird retained some of the reddish coloration on its mantle and a few pinkish tips to the flank feathers.

The other bird is like a good number ( 10 - 12 a season?) I see here in Connecticut. I have one at my feeder now. I do not really know its point of origin but am left wondering why we would get so many if it were truly a "vagrant" from a western population. Can a number of J. h. hyemalis simply show these western-like traits? Is it due to past hybridization at a western contact point between hyemalis and an "Oregon" form ? Is it contact with mearnsi? Is it "Cassiar Junco" or a product of Cassiar X Eastern Slate-colored?

I am curious to see your responses to the next set which I will try to get up tomorrow.

Mark S. Szantyr, Connecticut (14 Jan 2004)

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From Steven Mlodinow:

Greetings All

Unfortunately, I am busy trying to scurry out of town, so I can't enter this discussion in the depth I would like to.

The bird the Mark S has photographed in hand is a dead ringer for 50% of the juncos we have here in western Washington, and I can't really see the reason to put it into the Cassiar's category. The bird on the ground is messier, as others have noted, and I'd think it more likely what I'd consider as a possible Cassiar's. We see birds with some regularity here that look just like Sibley's depiction of Cassiar's, and it is my understanding that this is how adult males of that population typically look.

Cheers Steven Mlodinow, Washington (14 Jan 2004)

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From Mark Szantyr:

As promised, I have loaded three more junco photos from the east, Connecticut to be exact. for your consideration. This is fascinating to me and I hope will begin some considered discussion into what we know of this confusing group. The images are at http://www.surfbirds.com and follow the links to the North American Stop Press page.

Mark S. Szantyr, Connecticut (15 Jan 2004)

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From Matthew Kenne:

I was a little stunned by this statement earlier from Ted Floyd: "At the Powdermill banding station, for example, some 10-15% of the juncos (about 35,000 juncos in total) have been Cassiar." Do other banding operations in the East have anything similar to these results? I've always felt that the Cassiar Junco was the source of most of the western/Oregon-looking juncos we see in Iowa, but we don't see anything like a 10 % ratio with hyemalis. It's my understanding that the best determining factor in separating cismontanus from look-alike Oregons is that the Oregon had solid reddish coloration in the sides, while cismontanus has a mixture of reddish and grayish. Do those of you who have many cismontanus in the field find this mixture of color so difficult to see on a bird standing away from you with its feathers nicely layered that it's easy to miss? Is this actually the method used by banders in applying names to these birds or are there other features to look for? I'd really like to know what criteria they use at Powdermill for finding cismontaus among the hordes of imm/female brownish hyemalis birds.

Matthew Kenne, Iowa (15 Jan 2004)

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From Shai Mitra:

Hi all,

The recent junco thread in this forum comes just as I've just received several queries about the Fire Island, NY junco featured in the latest issue of North American Birds (Vol. 57, pp. 292-304). As my comments regarding that particular bird intersect with a number of more general issues raised in the recent junco thread, I thought a message to this forum would be most efficient and appropriate.

Several people have noted that in published photos, the Fire Island bird does not match their conception of Pink-sided Junco (mearnsi). Part of this is surely related to how the bird looked in life vs. how the photos look; another issue is the ongoing uncertainty and disagreement concerning how mearnsi (and other junco taxa) are expected to look.

> the start, the several photos give varying impressions. This is partly because of poor reproduction in NAB, and partly because my original prints were not very good. The task of lining a bird up with a particular taxon is bound to be difficult when the various photos seem to show different birds! That is where in-hand descriptions can be very helpful.

Certainly, the most striking thing about this bird in life was the pale, bluish-gray head with really black lores. This suite of features is not well-shown in any single photo, but can be pieced together from the various shots. That is, the paleness is suggested in one, the blue tone is suggested in some, and the loral feather tract is visible in the bottom-right shot, etc. This head pattern was strikingly different from those of the oddly colored female-type juncos we sometimes see in the East—and it impressed me immediately, even through the mesh of a bee-keeper's hat, before the rest of the plumage was really visible. Odd-looking, brown-flanked juncos in the East (call them what you will) almost always show darker, richer, browner heads that are relatively uniform across the lores, and that blend with brownish tones on the back, bib, and flanks. Similarly, female Pacific Oregon Juncos generally show darker, browner, and more uniform faces (although in these birds, contrast is usually evident between the head on the one hand, and the back and flanks on the other).

To my eye, Mark Szantyr's photo of the junco on the ground falls into the zone of messy brown eastern juncos. In contrast, the hand-held bird looks very unlike hyemalis to me, and also subtly unlike Pacific races of Oregon Junco. To me, its head and bib show the distinctive grayish tones of the juncos breeding in nw Montana and northern Idaho--Miller's 'montanus, southern division' (which roughly corresponds to Pyle's shufeldti).

In my opinion, this population of juncos is even less well understood than those around the contact zone in Alberta. Miller strongly emphasizes a north-south division of interior montanus. He describes southern montanus as much paler-headed than northern montanus--approaching mearnsi, and in fact intergrading with mearnsi in central ID. My recollection is that Miller describes females of southern montanus as paler-headed than those of the coastal population to the west--termed shufeldti by Miller, and simillimus by Pyle. This was also my impression based on examination of specimens. In contrast, Pyle describes coastal females (his simillimus = Miller’s shufeldti) as paler-headed than interior females (Pyle’s shufeldti = Miller’s montanus). I believe the discrepancy arises from north-south variation within the interior taxon.

To summarize, Miller characterizes females of southern montanus as:

Miller also emphasizes intergradation along the north-south axis between the two divisions of montanus (hence their inclusion in a single taxon) and between southern montanus and mearnsi, whereas Pyle emphasizes intergradation between montanus (Pyle’s shufeldti) and the population to the west (Miller's shufeldti = Pyle’s simillimus).

My own examination of specimens at AMNH has yielded the following conclusions on this confusing topic:

Getting back to particulars, the gray head of Mark Szantyr's hand-held bird strikes me as indicative of a Rocky Mountain bird; the absence of loral contrast, the browner (rather than pinker) tone of the flanks, and the restricted extent of the flank pigmentation all indicate Miller's 'southern montanus.'

In contrast, the Fire Island bird was really blue-headed, showed strong loral contrast, and pinkish flank tones—all indicative of mearnsi. Yes, the extent of its flank color was much more restricted than that seen in many mearnsi, but as noted above—and as stated in the NAB paper—the preeminence recently accorded this character remains debatable. The 4 May Deerlodge, MT specimen depicted in the NAB article actually does fall in line with the various southern montanus and mearnsi specimens in terms of overall plumage aspect. The date, locality, and striking loral contrast (perhaps obscured to some extent in the reproduction) all indicate mearnsi, despite the restricted flank color. Quite a few winter-collected mearnsi specimens also fall into this phenotype cluster.

Are birds like these merely intergrades between montanus and mearnsi? This seems possible. On the other hand, isn't it equally or more likely that mearnsi shows more geographic variation than the recent id literature implies? I tend to lean this way, because, whereas flank width and even head color clearly vary among the specimens in question, loral contrast seems much more clearcut. Core mearnsi show it, core montanus lack it, and its presence/absence among difficult specimens seems both clear-cut and congruent with geographic expectations.

Finally, it must be acknowledged that in a group as complex as the juncos, it might be impossible to rule out really oddball scenarios, such as Gray-headed X mearnsi or Gray-headed X Oregon. But should this sort of thinking be invoked when simpler interpretations exist? To summarize, I think that some Pink-sided Juncos resemble some Oregon Juncos more closely than is generally appreciated, but that these are nevertheless distinguishable on the basis of loral pattern.

Cheers, Shai Mitra, New York (15 Jan 2004)

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From Ted Floyd:

Hello, birders.

I've been enjoying the thread on juncos that Mark Szantyr got started last week. I have some additional thoughts on the subject, first in the matter of the identification of some of the photos that have been posted, and second (and more extensively) in the matter of the taxonomy and nomenclature of the genus Junco (with special attention paid to the race cismontanus, "Cassiar Junco").

I. Identification.

* Mark Szantyr's Connecticut birds.

- Mark raises the possibility that his Bird #1 contains hyemalis (sensu stricto) or even mearnsi genes. That's possible. Juncos interbreed extensively in their broad areas of overlap, and it's basically impossible to say that such-and-such an individual is pure. I continue to feel, though, that Bird #1 is a good match to cismontanus. By the way, Mark implies that cismontanus is a "vagrant" to eastern North America, but I do not necessarily agree with this terminology. The banding data from Pennsylvania indicate that cismontanus is regular and fairly common in winter at least into the western Appalachians. Meanwhile, Steve Mlodinow has commented that Bird #1 is a perfect match for 50% of the juncos that he sees in western Washington, which surprises me. Presumably, any junco that constitutes 50% of the overall population of juncos in western Washington would have to be something in the oreganus Subspecies-group. But several of us have opined that Bird #1 belongs to cismontanus. So this one is still up in the air?

- Regarding his Bird #2, Mark has revealed that it magically transmogrified into a more-or-less "normal" female-type hyemalis later on in the winter. That's interesting, and it is not inconsistent with my earlier speculation that the bird, although messy, does not appear to fall outside the range of variation for hyemalis and/or cismontanus. My hunch, based on Mark's photograph and description of the eventual condition of the bird, is that Bird #2 still ought to be considered problematic. I doubt it's all hyemalis. Steve Mlodinow has chimed in on this bird, too, which he considers to be a good match for an adult male cismontanus. But I'm not sure it's a male, and it's certainly not a "good" male cismontanus.

* Kent Nickell's Iowa bird. My first (and abiding) reaction to this bird is that it belongs to oreganus (sensu lato). Of course, there are multiple races within the oreganus subspecies-group. Does anybody have an opinion as to which oreganus subspecies is involved here? My guess would be shufeldti (=montanus), given the slightly contrasting lores, the pale cinnamon-brown tones of many of the contour feathers, and, especially, its occurrence in Iowa.

* Paul Buckley and Shai Mitra's New York bird. In another posting to this thread, Ned Brinkley mentioned this individual; there are four photos of it on p. 303 of the current (vol. 57 no. 3) issue of North American Birds. My hunch is that this bird is not a Pink-sided Junco (mearnsi), contrary to Buckley and Mitra's conclusion. To be sure, it is not a "good" Pink-sided Junco, e.g., one of the things bopping around my Colorado feeder right now. Its lores are not contrasting enough, its throat is not pale enough, its flanks are not pink enough, and the center of its breast shows too much hood and too little pink bulge. Could it have some mearnsi in it? Sure. Is it all mearnsi? I doubt it. Could it be cismontanus instead? I think so. Check out the adult female that is depicted in the bottom-righthand corner of p. 501 of The Sibley Guide. Also, compare it with Mark Szantyr's Bird #1 or with Fig. 3 on p. 135 of Joe Neal's article in the April 2003 Birding. Note further that Buckley and Mitra's biometric data place their bird within the range of variation for cismontanus, according to the treatment in Pyle's guide.

II. Taxonomy and Nomenclature. I should probably start off with a question that we've all been dancing around but that nobody's addressed head-on: What is a Cassiar Junco (Junco hyemalis cismontanus)? Apparently, it is a *stable*, self-sustaining population of juncos that arose via hybridization between hyemalis (sensu stricto) Slate-colored Juncos and shufeldti (=montanus) Oregon Juncos. The key here is "stable". We're at the point now--in terms of evolutionary and ecological history--that parental populations of hyemalis (s.s.) and shufeldti no longer are required for maintenance of the cismontanus population. If, in a thought experiment, we were to rid the world of hyemalis (s.s.) and shufeldti juncos, *we would still have* the race cismontanus. This result is quite in contrast with, say, flicker hybrids, or Passerina hybrids, in which intermediate populations depend upon a current crop of parental types. And neither are we talking here about a "hybrid swarm" situation, in the manner of, say, Western, Glaucous-winged, and Olympic (i.e., Western X Glaucous-winged) Gull hybrids. Again, we have a stable, self-sustaining population that appears to have arisen, at some point in its evolutionary past, via hybridization between hyemalis (s.s.) and shufeldti. There's a complication here, of course: cismontanus presumably interbreeds with both hyemalis (s.s.) and shufeldti, even at the present time.

And there's a bigger complication: If we assume that cismontanus is a stable, self-sustaining population, then it is not a straightforward matter to banish the trinomial (cf. Alvaro Jaramillo's posting). Then again, given its recent, hybrid origin, cismontanus may not exhibit any of the apomorphies, or derived character states (i.e., evolutionary novelties), that are usually considered to be required for the establishment of a valid taxon. In this scenario, Alvaro's proposal is looking pretty good again! The problem, really, is that the methods of cladistics--and of taxonomy and nomenclature more generally--are not equipped to handle the very real and very vexing problem of hybridization. The "Conventional Wisdom" in the North American ornithological literature is simplistic. The CW, in a nutshell: hybrids are aberrations; they represent population instabilities, fringe events, and evolutionary cul-de-sacs. Even in instances of extensive hybridization (e.g., Olympic Gull, above), our paradigm is to organize our thinking in terms of the parental populations. Buckley and Mitra, in their recent NAB paper, lament the "diminished emphasis on the taxonomic implications of hybridization", and I join them in their lamentations.

The one thing we all seem to agree on is that the hyemalis (sensu lato) and oreganus (s.l.) subspecies-groups are not valid. At a basic level, the existence of a stable, intermediate taxon (viz., cismontanus) would suggest otherwise. At a more fundamental level, I can't imagine that Alden Miller, or the AOU, or anyone else has actually established apomorphies for the hyemalis (s.l.) and oreganus (s.l.) subspecies-groups. You could argue that such groups, while not meeting the criteria for recognition as actual subspecies-groups, nonetheless constitute useful heuristic tools for birders and biologists in the field. But I question even this perspective: cismontanus simply cannot be pigeon-holed into either group, whether from the viewpoint of field-based heuristics or museum-based phylogenies.

The one thing we all seem to agree on is that the hyemalis (sensu lato) and oreganus (s.l.) subspecies-groups are not valid. At a basic level, the existence of a stable, intermediate taxon (viz., cismontanus) would suggest otherwise. At a more fundamental level, I can't imagine that Alden Miller, or the AOU, or anyone else has actually established apomorphies for the hyemalis (s.l.) and oreganus (s.l.) subspecies-groups. You could argue that such groups, while not meeting the criteria for recognition as actual subspecies-groups, nonetheless constitute useful heuristic tools for birders and biologists in the field. But I question even this perspective: cismontanus simply cannot be pigeon-holed into either group, whether from the viewpoint of field-based heuristics or museum-based phylogenies.

(Incidentally, the preceding analysis is tantamount--is it not?--to a ringing endorsement of David Sibley's decision *not* to use trinomial epithets in his field guides. You should either go "all the way", as Pyle does; or you shouldn't mess with trinomials at all. Otherwise, you'll have lots of Virginians calling their juncos hyemalis, in the strict sense, which they are not. That's the danger of using the NGS Guide.)

(Incidentally, the preceding analysis is tantamount--is it not?--to a ringing endorsement of David Sibley's decision *not* to use trinomial epithets in his field guides. You should either go "all the way", as Pyle does; or you shouldn't mess with trinomials at all. Otherwise, you'll have lots of Virginians calling their juncos hyemalis, in the strict sense, which they are not. That's the danger of using the NGS Guide.)

It's pretty safe to say that the Yellow-eyed vs. Dark-eyed distinction is bogus. The situation is reminiscent of the old Northern Oriole conundrum. As far as we can tell, Northern Oriole is paraphyletic--which, from a Phylogenetic Species Concept perspective, spelled its doom. Either it involves one species (Northern + Hooded), or it involves all three (Baltimore, Bullock's, and Hooded). Same thing with the recent split of the Black-billed Magpie, by the way, with hudsonia ("our" Black-billed Magpie) apparently more closely related to Yellow-billed than to pica (s.s.; "their" Black-billed Magpie). Thus, we have either one species of junco (combine Yellow-eyed and Dark-eyed) or we have three or more. And here's where it gets yet messier: For a variety of well-established and well-justified reasons of taxonomic theory, if dorsalis is separate from Yellow-eyed, then dorsalis is separate from caniceps (s.s.). But caniceps (s.s.) is still a part of Dark-eyed Junco. Or wait. No. How can it be? It has affinities with dorsalis, which has affinities with Yellow-eyed--but they're all separate species... How on earth are we going to pigeon-hole all of these junco taxa??

Sorry, folks. We can't. The botanists and the bacteriologists and the entomologists are all, more or less, coming around to this realization. Traditional species concepts simply don't work for American juncos. It's messy, plain and simple. There's no perfect solution. Apomorphies show up in the wrong places; hybridization is rampant, yet there are stable and distinct populations; certain individuals respond to the "wrong" species' songs; and on and on and on. There is, however, I think, a least- imperfect solution: Split 'em all. That's right. All of 'em. The Biological Species Concept is irrelevant in the case of junco taxonomy. But the Phylogenetic Species Concept addresses the junco conundrum with only a moderate amount of difficulty. There really are such things-- biologically and philosophically--as "Red-backed Junco", "Yellow-eyed Junco", "Slate-colored Junco", "Carolina Junco", and even "Cassiar Junco", the little bugger that got this whole thing started.

We birders in the field are 95% of the way to a solution. We're doing just fine, as long as we use good, useful, heuristically-sensible terminology, such as "Cassiar Junco", or "Red-backed Junco", or "Yellow-eyed Junco". It's when we start to get all mixed up on matters of trinomialism and taxonomy that the troubles begin. Mind you, I am emphatically *not* advocating taxonomic naivete. Rather, I feel, as Alexander Pope did, and as I suspect David Sibley does, that "a little learning is a dangerous thing". We can't go around slapping trinomials--or even binomials--on everything. I do feel that, ultimately, we can make taxonomic sense out of the junco situation. But we ain't gonna accomplish it with an outdated, irrelevant, BSC-or-bust mentality that clearly is irrelevant to junco taxonomy. Before getting to work on the eighth edition of their Check-list, I think that the members of the Check- list Committee should acquaint themselves with the recent literature on hybridogenesis, lateral gene transfer, and maybe even neo-Lamarckianism. It's a messy, messy world out there, and juncos are probably just the tip of the iceberg.

Best, Ted Floyd, Colorado (15 Jan 2004)

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From Mark Lockwood:

I thought I would add a little information that applies to Ted's comments about Gray-headed Juncos. The Red-backed Junco (J. h. dorsalis) breeds in relatively small numbers in the Texas portion of the Guadalupe Mountains. Interestingly, there are four documented records of Yellow-eyed Junco from that range. Unfortunately three of these are sight records, but one includes a very nice series of photographs. This individual clearly shows a yellow iris, but seems to be more like a Red-backed in plumage features. In particular, the rufous on the wing coverts is greatly reduced from what I consider typical for a Yellow-eyed. I am not familiar with the range in variation of this feature on Yellow-eyed Juncos and this may in fact be within that range. I wish I had the photos at hand to refresh my memory, but they are in the Texas Photo Record File in College Station.

The closest breeding populations of Yellow-eyed Junco are in southwestern New Mexico and southern Coahuila. This species would seem to be an unlikely vagrant to the Guadalupe Mountains, particularly considering there are only three other records for Texas (two in Big Bend and one in El Paso). I have wondered, on more than one occasion, if these apparent Yellow-eyed Juncos were not actually "yellow-eyed" Red-backed Juncos. As Ted mentioned, the thoughts on the placement of this taxon have oscillated between the two currently defined species. If in fact (if I dare use that term) Red-backed Junco are more closely associated with Yellow-eyed, then there may be "rare variant" yellow-eyed individuals.

On the broader topic on Dark-eyed Junco is general, there is an interesting hodge-podge of juncos that winter in northwestern Texas and many defy identification (at least for me). There appears to be almost a continuum of variation between Cassiar, Pink-sided, and Oregon. I have opined that intergrades must fill in the gaps, but that is about as far as I have gone. This weekend I looked at a fairly large group of Pink-sided Juncos with a scattering of Oregons and one male Slate-colored. The majority were what I think of as typical Pink-sideds, but at the time I wondered if I would recognize a female Oregon considering the variation observed overall.

Mark Lockwood, Texas (15 Jan 2004)

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From Vic Fazio, via Ted Floyd:

Hello, birders.

Vic Fazio, who browses this list but is not a subscriber, asked that I forward along his thoughts on intergrade juncos. Vic's commentary is relevant to several of the questions that have arisen here recently, including the following: How common is Cassiar Junco in the East? What was the Fire Island junco? Are birders under-reporting hybrid and intergrades? ... But I'll let Vic do the "talking" ... Best, Ted Floyd

In the mid-80's, I was bander-in-charge of the field stations of the Long Point Bird Observatory. By this time, LPBO had quite the dossier on variation in juncos and banders were encouraged to examine that variation carefully. In 3 fall, and 1 spring migrations, I personally handled about 1600 juncos, plus 300 banded at the mainland headquarters feeders over the course of two winters. Without going into too much detail, I observed the following . . .

No birds could be assigned to "Oregon" Junco.

Indeed, in several years at LPBO, I identified only one bird in the field as an Oregon. I recall this disappointing result as it was much less than expected based upon the relative frequency with which the form is otherwise reported in the east. Especially, as within my first month of banding at LPBO in Sep. 1984 I had already managed to capture an Audubon's Warbler and a RS X YS Flicker.

We did see birds we could not assign to Slate-colored as well. These we considered intergrades. These intergrades were almost exclusively recorded in MIGRATION. I can't recall the exact number for the winter records but it was likely but 1 or 2. In migration, I recorded 12 and 14 % respectively in 1984 and 1985 as intergrades[*]. I was not at all surprised at the results at Powdermill.

[*]These are my personal observations and are not part of the official LPBO database as intergrades were not recognized as a valid data entry at the time.

There was one curiosity regarding the winter flock. It was then and only then I recorded the white-winged Slate-colored Junco. Adult male Slate-coloreds with some pale tips to the greater coverts are not particularly rare in winter on the shore of Lake Erie. Birds with obvious pale tips the full length of the greater coverts comprised 10% of the group at LPBO. Of these, 3 were broadly tipped in white as were the median coverts. Still, they were otherwise clearly Slate-coloreds (although not nearly as dark as that variant's portrayal in Jim Rising's book).

Subsequently, I have been keenly studying juncos at a feeder in Tiffin (NW) Ohio since the fall of 1994. This feeder is on something of a migratory path for the junco as both spring and fall peaks routinely exceed 70 birds visible at one moment. Variation is considerable. In Ohio, I have identified 17 Oregon Juncos since 1977 (12 males, 5 females[**]). I think I am somewhat conservative regarding the ID. For a photo of one example from the Tiffin study site see http://www.aves.net/rarities/ojncopic.htm

[**]All adults; I have deliberately shied away from classifying birds appearing similar to HY female Oregon Junco (as depicted in Sibley) and place these with intergrades. I also do not try to separate out my Oregons. I would only note that the bird photographed above looks little different from those I have watched nest building in Belligham WA.

My dates on these Oregons range from mid-Nov. through mid- April. There are more records mid-Nov to mid-Dec. and mid- March to mid-April than in between. In other words, close examination of migrant flocks is as productive as mid-winter feeder study. However, very few birders in Ohio look hard at juncos before the advent of the Christmas Bird Count period. (Keep in mind that those CBCs conducted at the first opportunity within the count period will overlap, here in N Ohio, with the tail end of the junco migration; and occasionally take place smack in the middle of it.)

I record intergrades in Tiffin as well. I have recorded as high as 1 in 8 birds (12%) as an intergrade but 7-8% is more typically noted. Again, intergrades are usually less frequent in mid-winter, although I had two among 16 birds on 2 January this year.

This latter observation begs a couple of questions. In the hand, are banders more likely to determine an intergrade? Afterall, RS X YS Flickers are virtually never reported in Ohio yet, that 1984 season LPBO recorded 11. Is the lower apparent frequency of intergrades in winter a contributing factor in the overzealous reporting of western forms? (e.g most of the Cassiars moving into the east winter in the SE perhaps the mtns of TN, NC, WV etc. and it is the exceptional few remaining northward to visit feeders of the more populous Midwest and NE that give rise to erroneous reports of Oregons; could this resolve Matt's dilemma?)

I appreciate that in all this that some may also be wondering what I call a Pink-sided Junco. Here too is an example from Ohio (a record published in print in Ohio Bird & Natural History but otherwise overlooked). This bird possesses the classic sweep of color across the chest, the bluish cast to the hood, strongly defined lores, restricted white on the belly, etc.

See http://www.aves.net/magazine/mearnsi.htm

I therefore read with great interest the explanation offered by Shaibal Mitra regarding the NAB photos. I would have labeled it one of my intergrades.

BTW, Kent Nickell's bird is very interesting. Not along the lines of the intergrades I see in Ohio. Looks like an Oregon with washed out sides.

Cheers Vic Fazio, Ohio (15 Jan 2004)

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From Kenn Kaufman:

Comments below inspired by the rather long post by Ted Floyd earlier today. I should preface this by saying that Ted has been doing an outstanding job as editor of Birding magazine, with absolutely superb content in every issue, and that I agree with most of the things he's written in the last couple of years. On the "Cassiar Junco" (J. h. cismontanus), however, I think he is making some major false assumptions and employing some faulty reasoning, and thus making the situation at once simpler and more complicated than it really is.

Ted wrote: I should probably start off with a question that we've all been dancing around but that nobody's addressed head-on: What is a Cassiar Junco (Junco hyemalis cismontanus)? Apparently, it is a *stable*, self-sustaining population of juncos that arose via hybridization between hyemalis (sensu stricto) Slate-colored Juncos and shufeldti (=montanus) Oregon Juncos. The key here is "stable".

So, what's the evidence that there's anything "stable" about cismontanus? I have less experience in the Jasper Natl Park area than Alvaro Jaramillo does (see his post of 1/13), but my observations there and in parts of northern BC are about the same -- you see juncos there in summer that range from fairly typical-looking J.h. shufeldti to fairly typical-looking J.h. hyemalis, and everything in between. The situation looks like what I think it really is: a broad zone of intergradation between those two subspecies. What I think of as a "typical" cismontanus is just a bird that looks exactly intermediate between J.h. hyemalis and J.h. shufeldti. On either side of that hypothetical "typical" cismontanus, you'll find birds that look more and more like either of the respective parent groups; and at their extremes, it will be impossible to say whether a bird is cismontanus or just a grayish shufeldti (or brownish hyemalis). Similarly, it would be impossible to draw a precise boundary for the breeding range of cismontanus, because it will fade gradually into shufeldti to the southwest and hyemalis to the northeast. I don't see anything special about "cismontanus," as compared to any other case where two distinctive subspecies meet in a broad zone of overlap and intergradation.

To continue: We're at the point now--in terms of evolutionary and ecological history--that parental populations of hyemalis (s.s.) and shufeldti no longer are required for maintenance of the cismontanus population. If, in a thought experiment, we were to rid the world of hyemalis (s.s.) and shufeldti juncos, *we would still have* the race cismontanus. This result is quite in contrast with, say, flicker hybrids, or Passerina hybrids, in which intermediate populations depend upon a current crop of parental types.

What's the evidence that flicker hybrids depend upon a current crop of parental types? There is a broad zone of intergradation between Red-shafted and Yellow-shafted Flickers (especially broad in western Canada, but significant on the western Great Plains in the U.S. as well). Within this zone, if one looks closely at flickers in summer, it's almost impossible to find one that looks totally "pure" for either parental type. The interbreeding appears to be totally random. (The same random interbreeding occurs between Red-shafted and Gilded, by the way, but the areas where they meet are very limited.) I haven't seen any studies that show that these intergrade populations are less successful at reproduction, so I don't see why they would need a current crop of parental types. If all the "pure" Yellow-shafteds and Red-shafteds suddenly disappeared, I don't think the flickers in the zone of intergradation would wither away -- I suspect they would go on cheerfully reproducing, and that they'd continue to be a highly variable lot for many, many generations. How is this different from the situation with "cismontanus"?

To continue: And neither are we talking here about a "hybrid swarm" situation, in the manner of, say, Western, Glaucous-winged, and Olympic (i.e., Western X Glaucous-winged) Gull hybrids.

Actually, I think it would be perfectly valid to refer to "cismontanus" as a hybrid swarm, except that the term "hybrid" is usually used for crosses between two species, while crosses between two subspecies are more usually called "intergrades."

Again, we have a stable, self-sustaining population that appears to have arisen, at some point in its evolutionary past, via hybridization between hyemalis (s.s.) and shufeldti. There's a complication here, of course: cismontanus presumably interbreeds with both hyemalis (s.s.) and shufeldti, even at the present time.

I don't think cismontanus is stable, and I don't think "self-sustaining" is a relevant term. And of course it still interbreeds with both J.h. hyemalis and J.h. shufeldti. There are no barriers to separate the breeding ranges of those forms.

Cutting out a lot of stuff with which I agreed completely (what's the fun of that?), I wanted to pick on one other thing.

Ted wrote: The obvious next thing to do is to ask about the validity of any of the Dark-eyed Junco subspecies-groups. And we do, indeed, have many problems on this broader front. Let's shift gears and go to Arizona, where, until recently, most birders were happy to contend with two species of breeding juncos: Yellow-eyed (phaeonotus) in the southeastern > mountains and Dark-eyed (hyemalis, s.l.) in the mountains elsewhere. If you wanted to get fancy, until recently, you would call the widespread juncos caniceps ("Gray-headed Junco"), and that's where the trouble would begin.

Having been actively involved in Arizona birding for a long time, I have to disagree with this. For at least the last three decades, Arizona birders have been well aware of the differences between caniceps and dorsalis juncos. The typical Gray-headed Junco, J.h. caniceps, is a common wintering bird in southern Arizona. The breeding bird, J.h. dorsalis ("Red-backed Junco"), is pointedly sedentary on its breeding range in the central part of the state and is a very rare visitor anywhere south of those ranges where it nests. Birders here have long regarded J.h. dorsalis as being intermediate between Yellow-eyed Junco and J.h. caniceps in appearance, voice, and habits; and while we don't have the wisdom to say what this means about the larger questions of specific status of various juncos, we certainly wouldn't refer to this bird as caniceps for short. Indeed, dorsalis (unlike J.h. cismontanus) actually is a stable population in most of its range, and it might have been proposed for species status already, except that there are some breeding populations (e.g., north rim of the Grand Canyon) that seem intermediate between dorsalis and caniceps.

Also: After all, how many birders-cum-pedants in the southern Appalachians realize that hyemalis (s.s.) is *not* their junco.

Well, I can tell you that the first time I went birding in the mountains of western North Carolina, about thirty years ago, I ran into birders who proudly pointed out that the local breeding juncos were J.h. carolinensis. And when I was back there last fall, local birders were talking about picking out the smaller, darker migrant juncos from the larger, paler local breeding subspecies. I think, Ted, that the average birder may have more awareness of these issues than you give them credit for.

And finally, by the same token: But we ain't gonna accomplish it with an outdated, irrelevant, BSC-or-bust mentality that clearly is irrelevant to junco taxonomy. Before getting to work on the eighth edition of their Check-list, I think that the members of the Check-list Committee should acquaint themselves with the recent literature on hybridogenesis, lateral gene transfer, and maybe even neo-Lamarckianism.

Okay, Ted, you make the point that you like the Phylogenetic Species Concept (PSC) more than the Biological Species Concept (BSC in the above quote). Many systematists would agree with you, and you can see the evidence for influence of the PSC in many recent decisions on bird classification. But it seems a bit flippant to suggest that members of the AOU Check-List Committee "should acquaint themselves with the recent literature..." It's my impression that these people are in fact very well versed in the most recent literature and most recent thinking on taxonomic matters, even if we don't always like their decisions on specific cases.

At any rate, I'm certainly enjoying the junco discussion. Thanks to Mark Szantyr for kicking it off and to Ted Floyd for kicking it into a higher gear.

Kenn Kaufman, Arizona (15 Jan 2004)

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From Steven Mlodinow:

Greetings, Sorry for any misapprehension I may have caused. I do not think bird number two looks like a male Cassiar's. I think the drawing in Sibley is correct for male Cassiar's. I have no idea what Mark S's bird number 2 is. I don't think the photo is sufficient to allow ID.

Cheers Steve Mlodinow (15 Jan 2004)

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From Steven Mlodinow:

Greetings All
God, I love juncos, but I'm flying out of town very early tomorrow AM. So, I apologize for my brief post. I try to look carefully at most juncos I come across in Washington. The Fire I. junco would not cause me to pause here. The color of the hood, the extent and color of the flanks, and darkness on the lores is matched by a great number of our wintering juncos which are, of course, most unlikely to be mearnsi.

Off to look at Baird's Juncos.

Cheers Steve Mlodinow (16 Jan 2004)

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From Mark Szantyr:

As requested, a better view of the red junco from Feb 2002. Again, over the course of the 4 or so months I observed this bird, it changed from this rich reddish coloration to a mostly typical, dull, pale female-type gray hyemalis coloration, retaining some red in the mantle and a few reddish tips to the a few flank feathers. The image is at http://www.surfbirds.com and follow the links to the North American Stop Press page.

In order to keep these images and comments available, Angus Wilson has kindly offered to establish a page dedicated to this conversation on his Ocean Wanderers website so stay tuned for details on this by Angus.

Mark Szantyr, Connecticut (16 Jan 2004)

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From Ted Floyd:

Hello, birders.
Many thanks to Kenn Kaufman for his lively and thoughtful commentary on the "Nasty Junco" complex. Much of what he has to say deserves either (1) a response or rebuttal of some sort or (2) a cheerful acknowledgment that he is correct. In other instances, though, I detect in Kenn’s posting attempts at (3) mind-reading (always a dangerous enterprise in this electronic medium) or (4) jumping to the defense of perceived slights against his friends (which were not intended). I'll try to confine (most of!) my responses to the actual matter of junco identification (cf. 1, 2). After all, we're not psychics (3), and we're all friends (cf. 4).

Without further ado and, again, with all due admiration for Kenn's liveliness and thoughtfulness:

On the "Cassiar Junco" (J. h. cismontanus), however, I think he is making some major false assumptions and employing some faulty reasoning, and thus making the situation at once simpler and more complicated than it really is.

(What can I say? I aim to cover all my bases!)

So, what's the evidence that there's anything "stable" about cismontanus?

The direct answer, I guess is: "Because Alden Miller says so." I realize his classic revision of the genus Junco is now 63 years old, but it still seems to be the standard.

A seemingly less-direct, but actually more compelling, answer is: "Because the AOU recognizes cismontanus as a valid subspecies." There are hard-and-fast standards for elevating a population to the status of 'subspecies'. For example, there are various criteria of parsimony, synapomorphy, and--especially, in the case of AOU deliberations, as I understand the situation-- diagnosability. Subspecies need to be discrete, diagnosable, and stable. Thus, the AOU, following Miller's lead, regards cismontanus as a stable subspecies.

Of course, Kenn and others would be quick to point out--and I'm going to join the fray right here at the outset!--that the AOU's judgment on cismontanus was criticized almost from the get-go. (The "get-go", of course, was in 1957, when the AOU published the famous fifth edition of its Check-list.) By the early 1960s, Allan Phillips (Birds of Arizona) had already ditched cismontanus, on the grounds that only 1 valid race of junco was to be found in central Canada from British Columbia all the way to Quebec. Other writers, e.g., William Behle (Birds of Utah), have followed Phillips in this regard. Peter Pyle, the most recent authority on North American subspecies, basically follows the AOU in the matter of cismontanus, by the way.

Going back to Kenn's original question (What's the evidence for population stability in cismontanus?), I think we'd have to say: The jury's still out on this one. To be sure, there are birds that can be identified in the field as intermediates, and I do favor Alvaro Jaramillo's proposal to continue to give them an English name (viz., Cassiar Junco) without necessarily recognizing their trinomial epithet.

English names are a wonderful thing, pedagogically speaking. When I find myself in pedagogical situations, I make it a point to use names such as "Cassiar Junco", or "Olympic Gull", or "Brewster's Warbler", or good ole "Flin" (the bander's name for flicker intergrades). We--all of us--are
really handy with names. Everybody's talking about these charming "Avostilts" that are starting to show up in the western U.S. But if we were forced to call them "F1 hybrid progeny of H. mexicanus X R. americana", we'd lose interest pretty quickly.

Here's what I propose: Let's rehabilitate and promote the use of the name "Cassiar Junco", and let's get try to do the field work and lab work to answer Kenn's question.

Next: What's the evidence that flicker hybrids depend upon a current crop of parental types?

That's a fair question. I imagine there is no evidence. (Really, how can there be? I intended my analysis as a thought experiment, for what that's worth.) Note, though, that, given the evidence (or lack thereof), the AOU's stance on flickers is inconsistent with its stance on juncos. With flickers, we have a diagnosable eastern/northern population (Yellow-shafted Flicker; auratus subspecies-group), a diagnosable western population (Red-shafted Flicker; cafer subspecies-group), and an intergrade population that doesn't have a scientific name. But if flickers were juncos, we would call the intergrades a subspecies (could we call them 'flinosus'? please?) within, say, the auratus subspecies-group. That's what has been done with cismontanus, assuming you agree with the viewpoint that cismontanus is intermediate between shufeldti and hyemalis (sensu stricto); in fact, *even without* the stability/instability bugaboo, the assignment to one or another subspecies-group of a clearly intermediate population is unjustified.

If all the "pure" Yellow-shafteds and Red-shafteds suddenly disappeared, I don't think the flickers in the zone of intergradation would wither away -- I suspect they would go on cheerfully reproducing, and that they'd continue to be a highly variable lot for many, many generations.  How is this different from the situation with "cismontanus"?

As you've portrayed it, there is no difference. And if you're right--which you may well be (more data are needed, I'm sure)--then that's another nail in the coffin of cismontanus, and, more importantly, in the coffin of junco subspecies--groups more broadly. Oh, and just to beat a dead horse: Even if we wind up trashing the scientific name "cismontanus", please, can we continue to call intergrades "Cassiar Juncos"? Names, as I continue to maintain, are a wonderful thing. Without names ("Brewster's", "Flin", "Avostilt", "Olympic", etc.), things don't get named--and, as a corollary, they don't get noticed, they don't get studied, they don't get cared for.

More: Actually, I think it would be perfectly valid to refer to "cismontanus" as a hybrid swarm,

Maybe. But note that the population geneticists employ some amount of quantitative discernment in their characterization of "hybrid swams". I'm pretty sure the quantitative work hasn't been done, in the case of Cassiar Junco.

except that the term "hybrid" is usually used for crosses between two species, while crosses between two subspecies are more usually called "intergrades."

That's an arbitrary distinction. In the broader (ecological and evolutionary) literature, "hybrid" is definitely used to denote a cross between any two *populations*. To be sure, the process and its consequences (from gametogenesis right on up to life-history evolution) are the same, whether we're talking about interspecific hybrids (phaenotus X hyemalis, anyone?), intergeneric hybrids (Junco X Zonotrichia), or intraspecific hybrids (shufeldti X hyemalis; shufeltdi X cismontanus; even cismontanus X cismontanus, I suppose!). Anyhow, I don't favor the distinction between "hybrids" and "intergrades".

[Note: The rest of this message deals increasingly less with juncos and more with perceived slights against Arizonans, Southerners, mom-and-pop birders, and the AOU. Proceed with caution. --TF]

Having been actively involved in Arizona birding for a long time, I have to disagree with this.  For at least the last three decades, Arizona birders have been well aware of the differences between caniceps and dorsalis juncos

They have been? Where were they getting their information? The third edition of Peterson's western field guide (published 14 years ago) made no mention of dorsalis. Neither did the legendary second edition of the National Geographic Guide (published 17 years ago). And neither did the second edition of the Golden Guide (published 21 years ago). True enough, in recent years, field guides (e.g., Sibley, subsequent editions of NGS, and that wonderful guide with the Scarlet Tanager onthe cover) have started to make mention of dorsalis and/or Red-backed Junco. And if you go way back, to the 1961 edition of Peterson's western guide, there's extensive treatment of dorsalis/Red- backed; what goes around comes around...

But, for the most part, I'm skeptical of the claim that the great masses of birders in Arizona would have been well aware of the differences between caniceps and dorsalis, these past 30 years. In my own experiences in Arizona (which are, of course, not nearly as extensive as Kenn's), what I've encountered, by and large, is hordes of twitchers, tourists, and tarts. Nothing wrong with any of the preceding, of course. But I'll never forget the day (it was 2 July 1992, to be exact) that Chuck Bernstein and I went looking for Brown-throated Wren and Thick-billed Parrot in the Chiricahuas. All we got was good-natured contempt (emphasis on good-natured) for looking at birds that were some combination of (a) not in the book, (b) never heard of, and (c) not countable. We all had a grand ole time, mind you; I can't imagine, though, that the subject of dorsalis vs. caniceps ever came up. And I consider that particular day to have been typical of my experiences with *typical* birders in Arizona. (Meanwhile, I'm sure that the literati have long been aware of the dorsalis/caniceps divide.)

Birders here have long regarded J.h. dorsalis as being intermediate between Yellow-eyed Junco and J.h. caniceps in appearance, voice, and habits; and while we don't have the wisdom to say what this means about the larger questions of specific status of various juncos, we certainly wouldn't refer to this bird as caniceps for short.

And neither would "you" call it dorsalis. Come on, man! Go to some campground in the mountains, and ask the birdwatchers what they call that cute little bird poking about in the leaves. You'll get "junco" or "snowbird", or maybe "Dark-eyed Junco", or maybe "Gray-headed Junco" or "Red-backed Junco" (these days, probably about interchangeably). But dorsalis? I doubt it. We're talking about some pretty refined argot here. Most birders do not glibly and casually toss about terminology such as "dorsalis".

Well, I can tell you that the first time I went birding in the mountains of western North Carolina, about thirty years ago, I ran into birders who proudly pointed out that the local breeding juncos were J.h. carolinensis. And when I was back there last fall, local birders were talking about picking out the smaller, darker migrant juncos from the larger, paler local breeding subspecies.

I was in the region (central Virginia) last fall, too, and I had a delightful conversation with some local birders. Yes, birders. People with binoculars, bird books, and bird feeders. Y'know what we talked about?--"Black Snow Birds". It was a great conversation, and I learned a few things. For every southern birder who proudly points out J. h. carolinensis, there are ten times as many who happily revel in their Black Snow Birds.

I think, Ted, that the average birder may have more awareness of these issues than you give them credit for.

Next week, I'm leading a group of ca. 40 birders on a morning hike in the foothills here. We're going to look at juncos. For most of them, I assure you, the distinction between "Slate-colored" and "Pink-sided" is going to be unfamiliar; for some of them, in fact, the distinction between "junco" and "little brown job in the weeds" may be novel. For all of them, I'm virtually positive, the distinction between J. h. carolinensis and its congeners in the hyemalis subspecies-group would carry no meaning whatsoever. These are "average" birders. For every one of you or me, there are 40 (probably more like 400) of them.

Okay, Ted, you make the point that you like the Phylogenetic Species Concept (PSC) more than the Biological Species Concept (BSC in the above quote).

Actually, I didn't say so. Truth be told, I favor an approach that acknowledges the strengths and weaknesses of *both*. (By the way, there are about 25 other species concepts out there.) What I don't favor is a BSC-or-bust mentality. By the same token, I don't favor a PSC-or-bust mentality.

But it seems a bit flippant to suggest that members of the AOU Check-List Committee "should acquaint themselves with the recent literature..."

!!

Everybody should be acquainted with the recent literature. There's so much of it, and it's so diverse, and it's so good. I could be wrong (although I doubt it), but I would imagine that each and every member of the AOU Check-list Committee would agree wholeheartedly with me in the matter of the advisability of getting familiar with the recent literature.

Thanks, Kenn, for the great commentary and provocative questions.

Meanwhile, what –IS- this weird little thing outside my window, with lots of rufous on the back, and grayish flanks, and contrasting lores ... And, oh, no! -- this big woodpecker thingie on the tree, with brown-and-blackstripes and spots, and red--or, yellow?--in the wings and tail ...

Best, Ted Floyd, Colorado (16 Jan 2004)

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From Chris Elphick:

Hi all,
As I've told Mark before I don't have a clue what any of his juncos are, but I do have a couple of broader comments that follow up on Ted's posts. Apologies if this takes things a bit off-topic.

First, I think that Ted's analogy between the Yellow-eyed/Dark-eyed pair and the oriole and magpie examples is a bit misleading. It is possible that there is a similar pattern of evolutionary relationships going on with the juncos that would require taxonomists to either lump 'em all or split 'em further. But this is only true if Yellow-eyed is nested within the various Dark-eyed forms when the "tree" (cladogram, to get technical) of relationships is drawn. And the fact that there is apparently a close relationship between Yellow-eyed and Red-backed is not sufficient to demonstrate that this is the case.

It is hard (at least for me) to explain why this is so without drawing diagrams, but let me try. Imagine, hypothetically, that the ancestral junco looked just like a Yellow-eyed. These juncos were separated into two mountain ranges, and we ended up with some that continued to look like Yellow-eyed and some that look like Red-backed. Then from the Red-backed-like ancestors, all of the other junco types arose. This would put Yellow-eyed "outside" the group (i.e., they branch off first) that includes Red-backed and all the subsequent progeny species. If this is what happened then it is quite unlike the oriole and magpie examples (where either the hooded oriole or yellow-billed magpie was nested within the braoder group). This is because what we currently call Dark-eyed Juncos would be a monophyletic group (i.e., a group that all have a common ancestor and that includes everything that derives from that ancestor .... sorry for drifting into the jargon). Under this scenario the current taxonomy, though perhaps not very satisfying, is not necessarily wrong.

Ted's scenario could be right - e.g. if the ancestral junco were Slate-colored like, and everything else, including Yellow-eyed arose from that - but until someone figures out the exact relationships between all the different taxa and gives us a good well-supported phylogenetic tree, it is hardto know what should be done.

Second, although I tend to agree with all that Kenn Kaufman has written about the specific hybridization case involving Cassiar's Junco, I think that Ted had a valid point in highlighting the fact that so-called "good" species can arise from hybridization events. I don't think Ted said this explicitly, but this is something that commonly happens in groups of organisms other than birds - in plants it is really quite common, and it seems to occur at least occasionally in some vertebrate groups (e.g., some salamaders). This is one of the reasons why nearly all taxonomists who work on things other than birds seems to have abandoned the Biological Species Concept long ago (indeed as far as I can tell, most non-avian taxonomists never adopted it).

There has even been some suggestion of hybrid origins in bird taxa that are generally considered to be valid species, although I don't remember any certain cases. (Pomarine Jaeger is one bird for which this has been suggested, although I believe the latest evidence suggest that the hybrid origin is not the best explanation for this species. I seem to remember that there is some evidence that at least one of Darwin's finches might have arisen as the result of hybridization too.)

So, even though the evidence for Cassiar's as a distinct taxonomic unit (rather than just a hybrid zone - "hybrid swarm", I think, is generally reserved for cases where more than 2 species are involved) is apparently weak, in principle a "species" (whatever that might be!) could arise in this way.

Hope this hasn't just muddied one complex discussion with another equally complex one.

Cheers, Chris Elphick (16 Jan 2004)
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From David Sibley:

I have a few observations to add to this very interesting junco discussion. First, my experience is that birds like the ones Mark has photographed in Connecticut are regular in the east - not quite up to the 10-15% that are reported at Powdermill, but maybe 1 or 2%. There are three (of various appearances) among about 50 juncos at my feeders right now in Massachusetts.

I have seen photos of quite a few good Oregon juncos in the east, but I have personally seen only one undoubted Oregon junco in the Atlantic states, and that one really stood out. Even so, it was a female and it could have been a "Cassiar" cismontanus or some sort of impure Oregon such as a cismontanus x "Oregon" intergrade. The birds that Mark has photographed and that I am seeing out my window now generally inspire a minute or two of "maybe it is, maybe it isn't" study, and do not really jump out as something different. I have even seen quite a few of these birds that showed a well-defined convex grayish hood on one side and not on the other! And while there is a complete range of variation through the numerous "brownish Slate-coloreds" up to the few "somewhat Oregon-like" birds Mark has photographed, the variation essentially stops there.

There seem to be no males to go along with these females. Admittedly female juncos migrate farther than males (citations in BNA account) but still one would think that a few males would get this far, or that a few cismontanus of the more Oregon-like type would come along.

My inclination is to say that many if not all of these birds in the east are simply variant Slate-coloreds. They may come from some distant part of the wide breeding range of J. h. hyemalis, but I have doubts that they are cismontanus. One of the most interesting observations in this whole thread is Mark's report that the reddish junco in his photos from Feb 2002 transformed through wear and/or molt into a more or less normal Slate-colored by spring. Granted, this bird is not showing an Oregon-like hood, so may not be comparable to the others, but it raises the question of how much of this appearance is simply a brown winter aspect.

Banding records show a strong eastward component to the migration of Slate-colored juncos from the northern Rockies and Alaska, so states like Minnesota or Iowa might be getting some small numbers of "real" cismontanus from the Canadian Rockies. One banded in southern Yukon near the range of cismontanus was recovered a few months later in Minnesota (Sinclair et al 2003 Birds of the Yukon Territory). But I find it hard to believe that the relatively small and variable population known as cismontanus could account for the hundreds of thousands of brownish birds that must be spread across the eastern states if they make up even 2% of all juncos here.

Interestingly, in the early 1990s I saw several singing male slate-colored juncos in Denali Park in Alaska that showed fairly well-defined dark hoods suggesting cismontanus. I have no other confirmation of cismontanus-like birds occurring outside of the slate-colored/Oregon contact zone. Obviously these are not "real" cismontanus since they are so far removed from the presumed active-gene-flow Oregon x Slate-colored contact zone in the Canadian Rockies, but perhaps there is a tendency for far western Slate-coloreds to look like this, or maybe it's just a rare anomaly throughout the range. Any other observations from Alaska or northwest Canada would be very helpful.

I don't have an opinion yet on the Fire Island junco. It doesn't look like a Pink-sided to me, but given the disagreement between the written description and the photos I'm also not ready to call it cismontanus. If it is cismontanus it is still a very significant record, since I don't think many of these winter birds are.

As Shai Mitra points out, there is well-documented and undisputed interbreeding between Oregon (montanus) and Pink-sided (mearnsi) which absolutely must be taken into account when identifying these juncos. In addition the variation within Pink-sided may be more extensive than we want to believe. The question remains, if Pink-sided is variable, to what extent is this the result of interbreeding? The bottom line is that anything other than the brightest, most typical Pink-sided has to be distinguished from the intergrades, and some percentage of "pink-sided-like" juncos cannot be named with precision.

On the other hand, precision at that level is not always required. For example in this case if we could narrow the ID down to some permutation of Pink-sided ancestry (not suggesting that it is), then from a biological perspective rather than a listing perspective it makes little difference whether the bird is a pure Pink-sided from Montana or a Pink-sided x Oregon intergrade from Idaho. Both are equally out of place in the east.

Shai Mitra wrote: flank width and even head color clearly vary among the specimens in question, loral contrast seems much more clearcut. ...somePink-sided Juncos resemble some Oregon Juncos more closely than is generally
appreciated, but that these are nevertheless distinguishable on the basis of loral pattern.

This whole argument seems to overlook the question of interbreeding. The fact that lores seemed either blackish or grayish with few or no birds showing intermediate loral color is no reason to assume that contrasting dark lores indicate a pure Pink-sided and rule out an intergrade. One possible mechanism for this is that loral pattern is inherited as a unit, like the face pattern of Golden-winged Warbler which shows up in Lawrence's but not Brewster's hybrids. In this way some intergrades would have contrasting lores and be classified in this example as drab Pink-sideds, while very similar intergrades with gray lores would be classified as Oregon. I'm not saying that this is happening in juncos, just that it could explain the observation that loral contrast is either present or absent, while flank and head color covers the whole range of variation between the two subspecies. In any case we can't simply use one field mark as an on/off switch for this identification.

David Sibley (16 Jan 2004)

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From David Sibley:

It seems that my message and Angus's crossed in the ether. I've just taken a look at the photos displayed at http://www.oceanwanderers.com/JuncoID2.html, and wanted to clarify that I'm not lumping all of these birds as Slate-colored variants. One of the birds shown, figures 7-10, does stand out in my opinion as being beyond the range of variation of Slate-colored. As Mark says it looks good for Oregon, although I think it's actually a good candidate for a "real" cismontanus. It's certainly a western junco.

And Kent Nickell's photo from Iowa looks like a classic Oregon.

David Sibley (16 Jan 2004)

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From Chris Elphick:

Hello again, After my last post, David Sibley emailed me to ask a follow-up question. In the interests of full disclosure I should point out that I am no systematist so I'm not sure if can answer the question very well, but David suggest I post to the group so here goes. With luck some of the folks out there who I know know a lot more about this than me can chip in and clarify/correct.

David Sibley wrote: Isn’t it possible that the junco is one widespread species that has been differentiated by selective pressures? So that the Canadian Rockies birds are simply responding to an intermediate set of pressures or conflicting pressures from east and west and are not necessarily as Ted describes, a secondary contact zone? It just struck me that the scenario Ted lays out with such certainty may not be true.

My response: I think that what David is suggesting in his first question is that, rather than the hypothetical scenario of sequentially splitting species that I described, there could have been a widespread ancestral junco, that was subdivided into many subgroups, which then diverged and began to differ. This seems very plausible. My scenario was not meant to describe what I (or anyone else) think might have happened - it was purely hypothetical and was just intended to show that Yellow-eyed Junco could have Red-backed as its closest relative and still be appropriately classified as being a different species.

With respect to the northern Rockies populations that David asks about next, I would guess that the scenario David proposes is plausible .... but to my mind probably not very likely. I really don't know the junco literature very well, but based on what I know about hybrid zones in general I would assume that the situation is probably much like Kenn Kaufman has suggested. (Incidentally, based on his comments I think that David leans in the same direction, see below.)

David Sibley wrote:If the Cassiar Junco is really a stable intermediate population, they are no different than many other named subspecies along clines. I can’t think of any great examples right now (maybe the chain of Pacific Coast Fox Sparrows getting progressively lighter as one moves north – we don’t call the middle groups hybrid swarms). On the other hand, I tend to side with the people who think Cassiar Junco is a hybrid swarm like the contact zone of Yellow-rumped warblers in the same area.

My response: Again, I think David's reasoning here is right, although the relatively abrupt change suggests to me that hybridization after secondary contact is more likely. I think the bottom line here though is that we really can't know how these different populations differentiated from each other - and consequently what classification makes most sense - without someone sorting out the relationships between taxa ... and my guess is that this can probably only be done using genetic markers. (As for Fox Sparrows, maybe we'd think differently about them if they start to be split up!)

Finally, on another note, I think that someone in the context of this debate (although maybe it was some other similar debate) suggested that the AOU needs to "sort out" the taxonomy of the group. In general, I would argue that this isn't exactly the responsibility of the AOU's committee. Really it is up to the research community to do the work to figure out what is going on, and then up to the AOU to evaluate the studies and make a decision that keeps us all on the same page when we talk about these critters. Until the research is published, the AOU committee can do little but wait. Thus, their conservative approach on this (and other) issues seems entirely appropriate to me.

I hope this all helps, though I fear I may just be creating confusion with my vaguely informed digressions (at least I didn't get into polyploidy, paraphyly, and synapomorphies like I was tempted to - now that would have been nasty!). If someone who really knows something about systematics can clear up any mistakes I've made I'd appreciate it.

Chris Elphick

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From Kenn Kaufman:

Once again, thanks to Ted Floyd for insightful comments. I want to respond to one question he raised, not in order to be argumentative, but rather because it's an important point (one that I have emphasized while teaching identification workshops in the past).

In an earlier post, I had written: Having been actively involved in Arizona birding for a long time, I have to disagree with this. For at least the last three decades, Arizona birders have been well aware of the differences between caniceps and dorsalis juncos.

And Ted responded by asking: They have been? Where were they getting their information?

That's easy enough to answer. They were getting their information from _The Birds of Arizona_ by Phillips, Marshall, and Monson (U of A Press, 1964). This book is now way out of date for status of many species in the state, but it remains a classic for discussions of geographic variation and of subspecies. Allan Phillips was at swords' points with the AOU Check-list Committee for most of his career (and he had some weird ideas about species definitions) but he was brilliant at understanding geographic variation within species, and there's a huge amount about that subject in _The Birds of Arizona_. All of the birders that I was running around with in the 1970s just devoured that book, referred to it as "The Bible," and memorized the detailed discussions of variation within Fox Sparrows, Savannah Sparrows, Orange-crowned Warblers, Hermit Thrushes, and scores of other birds, including juncos (and yes, when we talked about J.h. dorsalis in casual conversation, the term was used for it was "dorsalis"). Even though the book is now out of print, scholarly AZ birders still track down used copies and study the book assiduously to understand what is going on with the variation that we see.

This is a point that I've tried to make repeatedly while teaching identification courses: One of your best tools for bird identification is not necessarily a field guide, but a work on status and distribution of birds in your local area. And it's vastly more useful if it's one of those books that really goes into the status of subspecies. Fewer such books are being published these days, since more of our information is based on sightings and photos, and less is based on specimens, so subspecies are harder to determine. But birders who really want to understand what they're seeing will seek out books like _The Birds of Arizona_, or Godfrey's _The Birds of Canada_, or Phil Unitt's _Birds of San Diego County_, or Oberholser's _The Bird Life of Texas_, or any other book that goes into detail on subspecies. Such references can take us to a completely different level of understanding of what's going on.

I know that you already know all this, Ted, but it seemed like a good opportunity to emphasize this point for people on the list who are still earlier in the learning stage and trying to figure out what references to use.

Kenn Kaufman, Arizona

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From Matt Kenne:

David wrote: My inclination is to say that many if not all of these birds in the east are simply variant Slate-coloreds. They may come from some distant part of the wide breeding range of J. h. hyemalis, but I have doubts that they are cismontanus.

I'd like to get back to the identification of Cassiar juncos in the East. If I assume (watch out!) that the breeding density of hyemalis juncos across the vast expanses of Canada and Alaska average out to anything similar to the average breeding density of Cassiar Juncos in their little corner of Canada, I estimate that the Cassiar Juncos make up 2%, maybe 3%, of the total hyemalis population. The Cassiar pool is drained further by birds migrating down the west side of the Rockies to winter in the Southwest (winters CA-TX by Pyle), and the North Pacific Coast: "Most of the Slate-colored Juncos wintering in coastal British Columbia appear to be J.h. cismontanus." (Rising, 1996); and this from Steve Mlodinow on Frontiers on March 29, 2001:

"I generally agree with Tony, except that intermediates between SC and OR Juncos are not rare in Washington. In w. Washington, I'd say the ratio of OR to SC is about 200 to 1. The ratio of SC to apparent SC X OR is about 4:1. Since we have lots of juncos, I see apparent integrades fairly frequently. Most look like cismontanus, though some don't."

So cut the cismontanus population available to migrate to the east by whatever figure you like (1/4, 1/3, 1/2 ?) - the ratio becomes appreciably less.

Three options come to mind that would increase the ratio between these populations by 5 or 10-fold to the levels reported from some locations in the East:

1. A difference in the percentage of the total of each group that migrate to the east. This probably occurs in the Southwest, as Slate-colored Juncos are much less likely to end up there, while a larger percentage of cismontanus do winter in that region (I'm assuming again, I don't have a reference other than Pyle about cismontanus wintering in the SW). This scenario doesn't really work in the East, the main wintering area for Slate-colored Juncos.

2. A Harlan's Hawk-type of migration strategy for cismontanus, whereby most of the group migrates to a core wintering range within the regular wintering range of hyemalis, increasing the ratio between the two groups while on migration and in the final stopping area. This doesn't appear to be the case with cismontanus juncos, or at least there's nothing in the literature to suggest it. It will be shocking if anyone has data or observations that indicate a mass exodus of cismontanus juncos out of the northern Rockies that ends with the group primarily wintering in the mountains of the Southeastern US- or anywhere else in the East.

3. Observers are confusing scuzzy, brownish, imm/female hyemalis for intergrades and identifying more cismontanus than are actually at their location. There just doesn't seem to be enough cismontanus to account forreported numbers. Compare the thoughts of Tony Leukering on intergrades in CO from this forum on March 27, 2001: "I have occasion to band a large number of juncoes in the fall here in eastern Colorado, so have handled 10s to 100s of all five groups, with WWJU the rarest. Generally, the birds I catch are distinct and separable to subspecies group, with little apparent intergradation/hybridization, though I must admit that I have seen a few that have defied identification - mostly immature females (really brown, dingy things); these may very well have been the product of some hanky-panky. However, the percentage of all juncoes that are possible hybirds - or, at least that show intermediate characters or a mixing of characters - is very low. Since eastern CO is blessed with migrant and wintering juncoes from across the breeding map, the very low percentage of obvious intergrades/hybrids may suggest that these actually occur in very low percentages in the metapopulation; I don't think that these "tweeners" are avoiding us." with those of Peter Yaukey from the Louisiana Bird Listserv on Jan 11, 2001:

"I spent my Master's work examining interracial dominance relationships among juncos in the Boulder area (CO) and in the Black Hills of SD in the wintertime, back in '85-'87 (Journal of Biogeography, 21:359-368). As Paul indicated, all forms can often be found in a single flock in winter in the foothills of the Rockies. This having been said, there were perhaps 10% of the juncos in my flocks that I felt were too "in between" to be assignable to race in the field. These probably included a number of the "cismontanus" Slate-coloreds that Paul mentions, which are hard to pin down in the field."

Is this a difference of in-the-hand vs. in-the-field? Same location, different interpretation, but the 10%-15% cismontanus numbers in the East are hand-held determinations.

I found the Powdermill webpage that the "10%-15% cismontanus" reference comes from. http://www.westol.com/~banding/Pictorial_Highlights_111318_2001.htm

It has a photo of a "fairly typical immature female cismontanus" to discuss if you think it warrants. It also has a description of their identification criteria:

Because extreme individuals of cismontanus approach them in coloration, and many bird watchers in the East are unaware of the existence of this intergrade race, it is likely that cismontanus frequently are mistaken for Oregon Juncos. This is particularly true for female cismontanus, which,like Oregon Juncos, usually have reddish brown side coloration thatcontrasts markedly with the lower edge of their gray hood (unlike Oregon Juncos, however, there usually is little or no contrast in the coloration ofthe upper edge of their hood and back). Importantly, not all juncos showing browinsh side coloration are cismontanus (females of nominate hyemalis oftenhave their plumage, including their sides, heavily suffused with brown)--thekey feature distinguishing cismontanus females is a fairly sharp demarcation between browish sides and a gray hood. In male cismontanus, the sides are alighter shade of gray, like a typical Slate-colored Junco, while the hood is nearly black, like an Oregon.

A discussion of these criteria might shed some light. Do these fieldmarks identify cismontanus from hyemalis to a reasonable degree of certainty?

Good birding, Matthew Kenne, Iowa

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From Kenn Kaufman:

In an earlier post, I mentioned _The Birds of Arizona_ by Phillips, Marshall, and Monson, an excellent source on subspecies but now out of print. Ian Paulsen informs me that you can still find copies of this volume on the used book market for about $20.

While we're on this major subspecies jag. Here in North America we're handicapped by the fact that the last "official" treatment of subspecies is literally half a century out of date: the last time the AOU Check-List Committee dealt with subspecies at all was in the Fifth Edition, published in 1957. So much work has been done since then that the 1957 treatment can no longer be regarded as authoritative. So, while we have an "official" list of which full species are valid, we have no such source for subspecies.

But in the absence of anything "official" we have something that's extremely good. Peter Pyle's _Identification Guide to North American Birds, Part I_ (Slate Creek Press, 1997) treats all of the passerines and near-passerines (doves, woodpeckers, hummingbirds, etc.), and includes an analysis of subspecies for all of them. Pyle did an extraordinary amount of work on this, not only studying specimens himself but also doing an exhaustive search of everything published on the subject in the last several decades; he describes the differences among subspecies and subspecies groups, and gives geographic distribution for all of them. I've found some places where I would disagree with his interpretations, but regardless, this is the best thing out there. Anyone with an interest in subspecies will find this book a gold mine of authoritative information.

Remarkably, I still run into some birders who say they're interested in identification but who don't have this book. I've said this before: This is the most important work on bird I.D. published in North America in the last two decades. Nothing else comes close. (Usual disclaimer: I have no connection with Slate Creek Press, nor with the book, except as a heavy user. I look at Pyle more often than at all my other I.D. sources combined.)

Kenn Kaufman, Arizona (17 Jan 2004)
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From Alvaro Jaramillo:

Birders,
Man, I feel old. As this junco issue was going on I was thinking of this project I did on the group when I was an undergraduate in University "just a few years ago". Well I finally found that bit of work and it was not a few years ago, it was well over a decade ago, nearing two now! Holy crap...where did all this time go? But you folks are surely not interested in my personal issues....let's talk juncos.

I measured skeletons (back in the early Carboniferous period) of 203 individuals housed at the Royal Ontario Museum. I dropped some locations due to small sample sizes, but had hyemalis from Ontario, Alberta (Hinton); White-winged from South Dakota (Custor Co.); Pink-sided from Saskatchewan as well as Okanagan Falls, BC (seeing that now, these were likely not Pink-sided, and I do note that in the results); Oregon (WAshington, BC); Grey-headed from New Mexico (Lincoln Co) and Arizona (Cocomino Co), I think both of these pertain to caniceps; Yellow-eyed (Chiricahuas, Arizona); and Volcano Junco from Costa Rica. There were 15 individual measurements, and then I looked at the stuff using a multivariate analysis called Principal Components Analysis.

The results were roughly this:

1) Morphological variation broke down to two major factors, one being size, the other being a shape component where some populations had a relatively (relative to their size that is, so think of this as a shape difference with the size element subtracted out if that makes sense) large flight apparatus (Sternum length, Ulna length), deep wide bill, and small legs (tarsometatarsus, tibiotarsus, femur, hallux lengths).

2) There was a main clump of juncos including Yellow-eyed, Oregon, Slate-colored, Pink-sided.

3) there were three outliers not part of the group in 2 (above) -

a) Volcano Junco is huge, has small flight apparatus, large legs and small bill. It was really an outlier! Note that this species is intermediate between Juncos and Zonotrichia and has been suggested to link the two genera.

b) Within North America the big outlier was White-winged Junco. It is really big, has a large bill, large flight apparatus, but small legs. Not sure why a Junco that apparently has a moderate length migration would have such a developed flight apparatus. Also note that studying specimens another odd feature of White-winged is that its shows much more reduced plumage dichromatism than the other northern Juncos. I suggest that the large size and lack of strong dimorphism could be due to intense competition for territories in their restricted, and isolated breeding habitat (Black Hills).

c) The other outlier but to a minor degree was Grey-headed Junco. It was intermediate between the White-winged and the rest of the North American Junco cluster in terms of size and the shape component.

4) Within the main Junco clump Yellow-eyed Junco was only moderately different in shape and size from Dark-eyed Juncos. Much more close to them than Grey-headed Junco was!!

5) Within the main Junco clump there was a general east-west cline; eastern birds being larger and having more well developed flight apparatus, shorter legs, larger bills.

6) Within the main Junco clump, Pink-sided Junco did not fit the general Cline. It was larger, with more well developed flight apparatus, shorter legs, larger bill. Closer to Yellow-eyed Junco; and somewhat intermediate between it and Grey-headed.

This doesn't help much with figuring out these confusing juncos that have been posted but it does offer an entirely different perspective on their geographic variation. Remember I did not measure or even look at any plumage details to arrive at these conclusions, they were all based on skeletons. Now if you put this data together with plumage, and the hybridization records one big question comes up, and it is the same one that came up way back when I did this? Given the large differences in hybridization and geographic variation between these different entities, why were the lines drawn the way they were drawn? Why is a Yellow-eyed Junco different from a Dark-eyed Junco? Another point is that sure there is a lot of hybridization between the Oregon and Slate-colored Group (= cistmontanus), but almost none between Pink-sided and White-winged. Given White-wing's large difference in structural morphology, small and specialized distribution, lack of strong sexual dimorphism, few hybrid records, consistent plumage differences from the other forms etc I do see how one could provide a good argument to split out that taxon. I think that one of the concerns a split of this taxon would be that in plumage it looks so much like a Slate-colored, but then go and compare Yellow-eyed (palliatus) with Red-backed (dorsalis) plumage and we seem to be comfortable with that. I would not be surprised if an eventual phylogeny either shows that Grey-headed and Yellow-eyed Juncos are more closely related to each other than we think, or that plumage patterns have no relevance to relationships based on molecular data. This would be similar to what is published on the Orioles, plumage similarity as with Hooded and Altamira is not matched by their phylogenetic relationship based on rather robust trees from molecular data for example.

cheersAlvaro Jaramillo, California (18 Jan 2004
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From Gary Felton:

More than usual, I've been scrutinizing the juncos at our feeders since the junco discussion began. Out of the app. 40, or so juncos that are coming to the feeders on both sides of the house, only one qualifies as a possible candidate for a female cismontanus (based on the illustration in Sibley). Two other birds might fall into that category, but it would really be pushing the limits. The rest of the juncos are a typical mixed bag of male and female hyemalis with several containing varying degrees of brownish and reddish tones on the back, head, sides and wings. The lower edge of the hoods vary from straight-edged to concave to very irregular with one side lower than the other. I'm too far north in the Alleghenies, so there's no carolinensis present, or none that I can detect anyway.

Regardless, if the cismontanus candidate is in fact a cismontanus, a ratio of 1 to 40 birds doesn't seem to correlate with the Powdermill banding stats and George Hall's statement in West Virginia Birds in regard to wintering juncos; "The evidence from birds banded at Morgantown indicate that the northwestern subspecies cismontanus Dwight is of frequent occurence."

Either the junco flock here is non-typical in regards to the ratio of cismontanus to hyemalis, there's cismontanus out there with hyemalis-type hoods and other hyemalis like characteristics, that I'm misidentifying as hyemalis, or there's no cismontanus at all and the bird here is just an unusual hyemalis. I, like others have said, also find it odd that male cismontanus never seem to turn up (I don't recall anyone mentioning a male to female ratio of cismontanus in banding data).

After I acquired the Rising and Beadle book, I suspected that I was occasionally seeing cismontanus at the feeders in the wintertime, but with all things considered, I have to wonder now if they are just not very reddish hyemalis (even if they do match illustration 48p on plate 21 in the Rising/Beadle book) Rising/Beadle illustrate a first-winter female while Sibley's bird is an adult female. It's too bad an adult female cismontanus wasn't included in the Rising book. Does anyone happen to have a photograph of a first-winter female cismontanus?

After speaking to one bird bander in regards to distinguishing cismontanus from hyemalis, I have to wonder if there aren't some misconceptions floating around in regards to cismontanus wintering in the east. I'm behind on reading some of the junco posts, but from what I've read so far and from what I've seen of junco flocks here in 25 years, I'm tending towards, I haven't seen a cismontanus yet. Since it appears that some female hyemalis may be indistinguishable from some female cismontanus, can someone provide me with any conclusive marks (bill color, body shape, anything) to distinguish the two in the field, (what is the banding criteria for making that determination)?

Regarding subspecies sources, it may be seriously outdated, but I've always appreciated the extensive list of subspecies that Peterson included in his second edition.

Gary Felton, West Virginia (19 Jan 2004)

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From Paul Hess:

Hello, all. During the tremendously educational junco discussion, a few people asked what criteria the banders were using to call a junco "cismontanus" at the Powdermill Nature Reserve in western Pennsylvania. Belatedly, I asked them to explain. The reply follows, posted with author Bob Mulvihill's permission. Best regards, Paul Hess, Pennsylvania

For the most part, we use identification criteria detailed in Miller's (1941) classic work "Speciation in the Avian Genus Junco," to which I refer contemporary birders with a serious interest in the subject. Importantly, we use Miller's rather loose definition of the subspecies to include virtually all intermediate Oregon/Slate-colored individuals (a very small number is intermediate so strongly in the Oregon direction that we band them as "best-fit" Oregon Juncos). In his words:

"Since, genetically, and certainly phenotypically, birds of immediate hybrid origin are identical with those of relatively uniform breeding populations of cismontanus, all birds of this appearance must be designated by this name when on the wintering grounds or on migration." (Miller 1941:342-343)

In digesting the pages of descriptions of various hybrids between the old-named J. oreganus and J. hyemalis (i.e., including, but not limited to, the geographic race J. h. cismontanus) found in Miller (1941), the take home story (and this has been confirmed for us both with live birds at the banding station and with specimen material from Carnegie Museum by Section of Birds' Senior Curator Emeritus, Dr. Kenneth C. Parkes) is that in female cismontanus there tend to be brownish sides and, from Miller's (1941) description, a "constant [grayish] neck line." This gives a fairly sharp (Oregon-like) demarcation between the gray of the lower edge of the hood and the brown (sometimes mixed with light gray) of the sides. The definition of the upper hood and back, however, is never distinct in cismontanus as it would be in a true "Oregon" junco. In male cismontanus, there is no brown in the sides at all, but a discernable "constant" neck line nonetheless is evident between the very dark (almost black) lower hood and the lighter gray sides. It is the females of cismontanus, only, that have the potential to create confusion (and very frequently do) with regard to "Oregon" and/or "Pink-sided" juncos in the East. Male cismontanus strongly resemble nominate hyemalis, and it is only with a lot of experience (and close examination in good light) that some can confidently be acribed as cismontanus. Consequently, our banded sample of cismontanus birds is heavily biased (about 5 or 6:1) toward females.

There is a fairly good depiction of this difference between the sexes in cismontanus (although he does not refer to the race by name) in Sibley's guide, in a boxed inset at the bottom right of p.501, showing an adult male and adult female Slate-colored Junco from the Canadian Rocky Mountains. As Sibley says, "some [females are] indistinguishable from Oregon in this broad intergrade population." Birders should note, in particular, that the gray hood and brownish sides in female cismontanus (and to a less obvious extent in males) create a sharp point or corner, when the bird is viewed from the side or front, whereas in simply overall brownish female hyemalis, the hood and sides blend together in a smooth line, with no visible point or corner between them. Honestly, it is this feature, which we depend on most of all for spotting the one or two cismontanus among the ten or more hyemalis we band.

To set the record straight, although cismontanus can comprise up to 15% or more of some samples of our migrating or wintering juncos, and this is a conservative estimate of the maximum to be sure (i.e., given the difficulty of spotting the males on a busy day when we're processing hundreds of birds), 5% or less of our total sample of juncos (which is about 40,000) actually was judged at the time of banding to be cismontanus (i.e., based on comments in the original banding record).

Finally, the information in Bob Leberman's Birds of the Ligonier Valley on this matter is very good:

"There are several records...of so-called "Oregon Juncos," formerly considered to be a separate species, Junco oreganus. None have been collected in our area, but specimens from eklsewhere in the northeast generally have been referred to the stongly migratory subspecies Junco hyemalis (formerly oreganus) montanus of the norther Rocky Moutnians. Rather commonly seen in our area, especially in March, is another form that may sometimes constitute as much as 20% of the migrating or wintering population. These jucnos are a rather variable array of intermediates between the "Slate-colored" and the "Oregon" coloration. They may come either from areas where the two kinds of juncos meet and interbreed (which is why they are no longer considered separate species), or from an area in western Canada where a more or less stable population of intermediate juncos has been described as the subspecies J. h. cismontanus."

Robert S. Mulvihill, Powdermill Nature Reserve, Pennsylvania (21 Jan 2004)
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From Barb Beck:

This is a question that is bothering me and I do not have the necessary literature available. Are the older studies of "cismontanus" possibly limited by the availability of specimens over a sufficient area? I am sure the work is excellent but were the conclusions possibly somewhat shaped byspecimens available. In the 40's and 50's in California where I am from you could cover most regions by driving and hiking. In areas of Canada like Alberta this was not the case particularly during the breeding season for these birds. Roads were few and basically mud during this season - bridges just not there. For the most part you went where you could take a train (if available) , horse, walk or boat. Vast areas required very great effort to access during the breeding season if you did not have wings.

Incidentally - although the southern half of Alberta is now crisscrossed by highways and well maintained gravel roads, we are just getting spring and summer access (thanks to oil, gas and pulp mills) to a vast block of NE Alberta which is about a quarter the size of the province - and this province is big.

Barb Beck, Alberta (21 Jan 2004)

PS If you think some of the birds in Western Canada are nasty because they are meeting after being split by glaciers (sometimes as good species, sometimes as clearly subspecies and sometimes as something inbetween) - try to sort out our butterflies who were not only split east and west but had populations north of the glaciers and possibly contact with Asian species over a land bridge.

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From Matt Kenne:

Mr. Mulvihill wrote:
In digesting the pages of descriptions of various hybrids between the old-named J. oreganus and J. hyemalis (i.e., including, but not limited to, the geographic race J. h. cismontanus) found in Miller (1941), the take home story is that in female cismontanus there tend to be brownish sides and, from Miller's (1941) description, a "constant [grayish] neck line." This gives a fairly sharp (Oregon-like) demarcation between the gray of the lower edge of the hood and the brown (sometimes mixed with light gray) of the sides.

Everyone digests things a little differently. I can't draw the same conclusions when I consider Miller's statements from p. 332: "Thus slate sides are not dominant or at least do not appear with great frequency, because of interaction of sex factors or hormones in females." " ...but out of 32 females from the same region, 26 showed some buff. (6 birds then did not have buff in the sides- MK) The confluency of the head and sides is another matter, however. Occurrence of an even gradient in side and hood color is essentially as frequent in females as in males in groups of dual affinity."

In male cismontanus, there is no brown in the sides at all, but a discernable "constant" neck line nonetheless is evident between the very dark (almost black) lower hood and the lighter gray sides.

This is plainly not the case, on p. 330 it states: "In males, obvious hybrids show nearly every degree of mixture of slate and brown. The two colors are segregated into different areas of the feather vanes. Then anterior part of the side area is more prone to slate color, the posterior region to buff and brown. No occurrence has been found of an anterior section browner or buffier than the posterior section."

This is thought to be a trait from the hyemalis influence where a gradient of slate color passes from the head to the sides.

On p. 331 it states: "The independence of the gradient and pigmentation factors is shown by hybrids in which the sides are pure gray yet distinctly marked off from the dark slate of the hood, and by others with buff or brown sides which anteriorly have some slate that grades into the hood." That's grading into the hood without a hood line.

On p. 332 it states: "Out of 41 males of the Cassiar area, 7 showed some buff in the sides," Not all buff sides, possibly indicating Oregon parental types, but some buff in the sides. While a preponderance of males have hyemalis-like all-gray sides, it is clear that a portion does not.

The definition of the upper hood and back, however, is never distinct in cismontanus as it would be in a true "Oregon" junco.

I don't have the complete Miller reference here, just select copied pages, but in a margin I have noted "Cismontanus from hyemalis by hindneck line in 80%-95%." I believe it's from somewhere in Miller, but I'm no longer sure if it meant that 80%-95% of the specimens and observations from the Cassiar breeding area have this mark, indicating that up to 20% of the birds are "pure" hyemalis types; or if it means that up to 20% of the Cassiar birds have to be differentiated by other means, such as hood/side line or mixed gray and brownish in the sides. A long paragraph on p. 332 describes the variation in the contrast of the hindneck line from lineless in hyemalis, through an increasing degree of darkness on the head in cismontanus "...over the neck to a definite line which, with greater intensity of black, becomes more and more apparent." As both montanus and hyemalis can overlap in the depth of black tone on the head (dull black or dark slate), a hybrid at the Oregon-end of the spectrum would have head-back contrast (and line) identical to parental Oregon forms.

Birders should note, in particular, that the gray hood and brownish sides in female cismontanus (and to a less obvious extent in males) create a sharp point or corner, when the bird is viewed from the side or front, whereas in simply overall brownish female hyemalis, the hood and sides blend together in a smooth line, with no visible point or corner between them. Honestly, it is this feature, which we depend on most of all for spotting the one or two cismontanus among the ten or more hyemalis we band.

The first mark I look for in Iowa for attempting to identify a Cassiar in the field is the presence of a hindneck line. This line conclusively shows the effects of Oregon influence. Consider the bird on the Powdermill webpage http://www.westol.com/~banding/Pictorial_Highlights_111318_2001.htm
There is no obvious hindneck line in these shots. Lacking this line, it had better have a very obvious, sharp demarcation line between brownish sides and a gray hood to consider it an intergrade over a hyemalis with brownish wash, and the Powdermill bird doesn't qualify in my opinion. Perhaps if I had the bird in hand? Both males and females (according to my reading of Miller, above) occasionally lack the dividing line between the hood and the side. At this point in the field, I'm left with trying to decide if there is a mix of brownish and grayish basally in the feathering of the sides- not just a wash of brownish through the top. I've never been successful in this to my satisfaction.

Finally, the information in Bob Leberman's Birds of the Ligonier Valley on this matter is very good: Rather commonly seen in our area, especially in March, is another form that may sometimes constitute as much as 20% of the migrating or wintering population.

"especially in March" is important in this observation. I've always noted an increase of "funny-looking" brownish juncos as the spring progressed. In discussing hyemalis on p. 315, Miller states: "Wear always tends to purify the gray and slate of the hood and to make it slightly darker, for the bases of the vanes are heavily pigmented." This would tend to turn those brownish hyemalis juncos into more hooded, intergrade/Oregon-like birds. It's kind of like having a frustrating flock of gulls in my own backyard!

Matthew Kenn
, Iowa (21 Jan 2004)

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From Tony Leukering:

Hi all: One more response on this thread. Well, actually, two, but both in this message.

1) Alvaro's results are quite interesting, but more and more what I have come to expect in juncos (and bird biology, in general) -- the more things look similar, the more unlike their whatever (DNA, structure, voice, etc.; there may be a good reason for this involved with assortative mating of different taxa). Of course, there are taxa that look similar that are actually closely related, but.... Adding to Alvaro's comments about Pink-sided and White-winged (and forgive me if I have already mentioned these items in a previous post, one does get old and forgetful), Christopher L. Wood (not the Chris Wood in WA) and I have worked a fair bit on juncos here in Colorado and we find that Pink-sided is the taxon most similar, in many respects, to White-winged of all the juncos that we get here. Pink-sided and White-winged share the pale gray coloration, the black loral patch, and the extensive white in the tail. In addition, Pink-sided's call note is more similar to that of White-winged (which is fairly readily distinghishable) than are other "Dark-eyed" forms. Finally, Pink-sided is among the largest of juncos.

Alvaro's data suggest that junco taxonomy is not simple (what a surprise, huh?). In the northern juncos (SC, PS, WW, OR), Slate-coloreds get smaller from east to west (and the ones I deal with here in CO are pretty small) and Oregon is pretty small. Somewhat between those two (but more southerly) is Pink-sided, which is considerably bigger than either. Pink-sided is the closest breeding taxon to breeding White-winged which is, obviously, quite large. Now, head down the spine of the Rockies and one gets all large forms (Gray-headed then Red-backed then Yellow-eyed). Slate-colored and Oregon have the least amount of white in the tail. Pink-sided and White-winged have the most. The three red-backed taxa are intermediate.

The geology of the Rockies creates relatively discontinuous breeding ranges for the various junco taxa. The red-backed taxa (specifically, Gray-headed) seem not to be in contact with any of the "northern" juncos, though the Bighorn Mts in ne WY provide a possible contact point between Gray-headed and Pink-sided and (possibly) White-winged. (I don't know anything about possible contact zones in the Great Basin, an area with which I am almost wholly unfamiliar; Sibley does not indicate such a contact zone). However, all forms are migratory to greater or lesser extent, so it is certainly likely that individual birds find themselves in the wrong place, but I don't know where one finds direct contact of Pink-sided, White-winged, and/or Gray-headed on a widespread basis. I realize that the northernmost Rockies in the US may be continuous enough to support continuous contact of Oregon and Pink-sided over some area. Also that Alberta is a mess. (I don't see how anyone can remain sane birding in Alberta -- all the horrible contact zones are there: sapsuckers, "Solitary" vireos, juncos - ugh! They obviously make Alberta birders of more stalwart stuff than your average bear (to paraphrase Yogi.))

So, this is just a long and drawn out way to say, "HELP; we need help! Are you hybridizers of DNA listening? Please sort this mess out for us"

2) In regarding the apparent differing views of Colorado wintering juncos presented by Peter Yaukey and me (quoted in Matt Kenne's post; and if Peter is reading this, "hi"), I am comfortable calling a goodly number of these "apparently" intermediate birds "cismontanus," thus they are Slate-coloreds (per A.O.U.); Peter may not have been. This could readily account for the discrepancy, since a fairly largish number of Slate-coloreds that I see here (perhaps as much as 10-15%) are what I have been calling "cismontanus" (birds with the colors of Slate-colored, but with the pattern (distinct hood) of Oregon).

Well, how about a third.

Regardless of one's ideas on taxonomy (one junco vs. two junos vs. five juncos), I still do not understand why Pink-sided was so slighted as to be considered merely yet another form of "Oregon Junco" (sensu lato) when it is at least as distinct, in my opinion, as many of the other subspecies groups.

Sincerely, Tony Leukering, Colorado (22 Jan 2004)

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