MOUTH LINING: Lanyon (1961) said that this "is the most practical and most reliable morphological criterion for the specific identification of fresh specimens of this complex." Sounds promising! He "found no evidence that the character varies geographically or sexually in either species. In my series of 50 specimens of known voice, there was 100 per cent agreement between the specific determination based solely on color of mouth linings and those based on certain diagnostic vocal patterns." Nutting's has "orange-colored mouth linings." All fresh Ash-throated specimens had "'flesh-colored' mouth linings ('pale ochraceous buff' of Ridgway 1912)." He concludes by noting that Nutting's groups with Great Crested and Dusky-capped on mouth lining color, while Ash-throated groups with Brown-crested. Lanyon's mouth-lining character has been cited over and over in the literature as diagnostic.

Just reading the above, however, raises some questions. Does his "series of 50 specimens of known voice" imply there were a maximum of just 25 of each species? If so, that is a fairly small series compared with the over 400 specimens he used to develop his tail pattern key. Secondly, although he found no geographic or sexual differences, he doesn't mention age-related distinctions at all. His study was primarily focused on the question: "is Nutting's a good species?" (he found it was). But it is clear he focused on birds on territory — in other words, the evidence suggests very strongly that his 50 specimens were all adults on the breeding grounds. Thus he does not answer the question: is there age-related variation in mouth lining color?

This is an important question. Cardiff & Dittmann (2000), writing about Brown-crested Flycatcher and discussing its separation from Great Crested, describe mouth linings. As we have seen above, Nutting's is said to group with Great Crested and have orange mouth linings, while Ash-throated groups with Brown-crested with flesh-colored linings. Cardiff & Dittmann write that "not all [Great Crested] have bright orange-yellow [mouth linings]; LSU specimen data (described as flesh, dull-yellow, and dull yellowish-tan; these atypical mouth colors may pertain to first-year birds, but more data needed);" [emphasis added].

If it is possible that the bright orange mouth linings of Great Crested can be dull yellow or flesh-colored in first-year birds, is it possible that the same may be true in Nutting's? Given this question — for which there is no data — I don't see how anyone can conclude that mouth lining color will diagnostically identify the Santa Cruz bird, since we know for a fact that it is a first-winter bird.

Further, mouth lining color variation also goes the other direction. There are specimens of Ash-throated Flycatcher whose data tags state the mouth lining was "orange" (Bowers & Dunning 1987, cited with approval by Cardiff & Dittmann 2002).

Bowers & Dunning (1987) published a widely-read paper in American Birds on a bird they had captured and banded in Santa Cruz Co., Arizona, on 15 July 1985 (not to be confused with Santa Cruz Co., California; all reference to "the Santa Cruz bird" in these pages is to the California bird). They identified it as a Nutting's, and published both photos of it and comparative material. The bird was heavily worn and called only once upon extraction from the net; the identification as Nutting's relied heavily on the presence of orange mouth linings. This identification is now considered controversial (Cardiff & Dittmann 2002); there is an on-line statement that the Arizona Bird Records Committee thinks it likely a worn Ash-throated (although they have never reviewed the record). If it is an Ash-throated, this is another orange-mouthed Ash-throat beyond the specimen cited by Bowers & Dunning (1987).

I noticed some interesting points in reviewing the published photos; I have scanned the relevant pages and reproduce R.W. Bower's photos here (fair use doctrine; these shots are published at Amer. Birds 41: 8 & 9).

The first shot (near right) is an Ash-throated that is stated to have "pink" mouth lining. The second (far right) is the mystery bird, considered Nutting's in the article, stated to have "orange" mouth lining. I don't see anything that is nearly that simple. First, the tongues of both birds are bright pink. Second, the color inside the lower mandible on both birds is a really dull straw color. The "mouth lining" itself must refer to the lining deeper in the mouth and (maybe) inside the upper mandible, but it cannot mean the tongue or the inside of the lower mandible.

The key point on mouth lining is the color inside the actual mouth itself, including (perhaps) the "roof" (upper mandible lining) but excluding the lower mandible lining, and (probably) excluding the tongue. Below is a series of four video-caps (© Joel Weintraub) of the Orange Co., southern California, Nutting's:

If the key point is the actual color of the inside of the mouth and upper mandible only, what colors should we see? As discussed above with reference to Lanyon (1961), Nutting's is orange while Ash-throated has "'flesh-colored' mouth linings ('pale ochraceous buff' of Ridgway 1912)." Cardiff & Dittmann (2002) describe Ash-throated mouth lining as "usually flesh-colored, occasionally yellowish flesh to dull yellowish, rarely orange." Please note the absence of any reference to pink, let alone bright pink, by any of these authorities. The only reference to "pink" in Ash-throated that I find is in Bowers & Dunning (1987), and I suspect it is a reference to the tongue. Look at their in-hand photo of their undeniable Ash-throated, reproduced above. The tongue is pink, the bottom of the lower mandible is a really dull straw, but there is a tiny bit of the mouth lining itself shown just inside the gape and right of the tongue. It looks dull yellowish to me, and that color is consistent with the literature descriptions. You can go check up on "pale ochraceous buff" yourself at an online color chart; as shown there, it is the equivalent of pale yellowish-pink. It is rather like the color of the inside of the lower mandible on the in-hand pictures, above. It is sort-of like the color of the back of your hand if you are Caucasian and not tanned or sunburnt. It is not anything like bright pink. So we want to see if the Santa Cruz bird has a dull mouth-lining that is flesh-colored or pale yellowish-pink (=flesh), or a bright orange mouth lining that can look bright pink in some views (e.g., the Orange County bird, left hand shot above).

To help this analysis, I have used the PhotoShop color picker on the four shots of the Orange County bird, and then created a box with that color in the lower left of each photo, and then have done the same with the photos I can find of the Santa Cruz bird. I try to choose the color in the very center of splash of color and away from edges or shadows:

Orange Co. Nutting's [photos © Joel Weintraub]
Santa Cruz bird [photo near right © Cindy Cummings; far right © Peter LaTourrette]		No photos show this same angle but described as bright orange but most observers	No photos show this same angle but described as bright orange but most observers

So that brings us to the leftmost photo in each row. To my eye, both photos looked to have bright pink mouth linings deep inside the mouth and inside the upper mandible. I don't think the tongue is in the way. Yet the "objective" color picture shows those colors to be a dark maroon! I think this means that the actual color is somewhat shaded in these shots and our eyes automatically adapt for that shading to see a bright pink. I added a box that the color picker chose as the color of the primary edges to test Cindy Cumming's photo, and it looks "in the ballpark" for the right color, although tending a bit too brown. There is a remarkable similarity in the tone of color chosen by the color picker between the lefthand shots of both the Orange Co. and Santa Cruz birds. We are missing photos of the Santa Cruz bird with the proper angle of direct light, but, given the experience with the southern California photos, I would not be at all surprised to find they would show bright orange. In any event, I hope we can all agree that Cindy Cumming's photo shows a bird with brightly colored mouth linings, not dull colored.

Incidentally, most observers of the Santa Cruz bird describe the mouth lining as orange. But what are they describing? My view was facing the bird as it ate an olive, I believe the tongue was below the olive, and I believe I described the color of the upper mandible lining, just as shown in the photos of the Orange Co. bird, center shots.

Is there a possibility that those who have described a "flesh-colored" mouth lining were looking at the lining of the lower mandible, as in the bottom row, far right, photo above? You can determine whether that is a possibility, and whether you want to give any weight to the descriptions of "orange" mouth lining from various other observers.
 

VOCALIZATIONS: In reviewing the literature, I notice three interesting points about vocalizations among the Myiarchus flycatchers in the group that is comprised of Nutting's, Ash-throated, Brown-crested, and Great Crested Flycatchers:

• First, the standard North American field guides (e.g., National Geographic 1999) and the birder i.d. literature (e.g., Howell & Webb 1994) emphasize the single diagnostic vocalizations and don't mention the other calls given by the species. As we shall see below, each species in this set has at least four standard calls. In his brief i.d. summary, Lanyon (1997) says the diagnostic calls are: wheep! for Great Crested, WHAY-burg for Brown-crested, ka-BRICK for Ash-throated, and (in Lanyon 1961), a clear penetrating whistle peer for Nutting's. National Geographic (1999) renders these as: a loud, whistled wheep in Great Crested, a rolling whit-will-do in Brown-crested, a rough prrrt and a burry ka-brick in Ash-throated, and a rather sharp wheep in Nutting's. Cardiff & Dittmann (2000, 2002) say the key Brown-crested call is a diagnostic rasp. A chart in Howell & Webb (1994) says the call notes are: rising wheep in Great Crested, a two-syllable h-whick in Brown-crested, a quiet pip in Ash-throated, and an emphatic wheek! in Nutting's. About the only things that seem sure in this review of vocalizations is that Great Crested gives a rising wheep and there is a ka-brick (emphasis on second syllable) in Ash-throated. Other than that, even these experts don't seem to completely agree on what the diagnostic vocalizations are.
• The second interesting point has to do with the evolution of what call of Nutting's is considered diagnostic. As will be discussed below, Lanyon's (1961) major study found four basic Nutting's vocalizations. The one he considered most diagnostic was the clear, penetrating whistle peer, sometimes lasting longer peeer that was suggestive of the standard call of Dusky-capped Flycatcher but "usually at a higher frequency" and "more 'piercing', less 'sad' or plaintive." Lanyon tape-recorded this vocalization in all populations of Nutting's that he studied. By the time of Bowers & Dunning (1987) they said the common call of Nutting's was "a smooth, rising, single-syllable wheep or cheep." Lanyon (1997) kept his description as peeer for Nutting's, and so did Dittmann & Cardiff (2000), but Howell & Webb (1994) used "an emphatic wheek", sometimes rendered as wheep (Howell & Webb 1995), and, as we have seen, National Geographic called it a "rather sharp wheep." Therefore, in short, Lanyon's peeer evolved into a Great Crest-like wheep! over the course of the decades, and the conventional wisdom is that Nutting's gives a call similar to Great Crested. This is a curious evolution, and the conventional wisdom may not be entirely accurate.
• Dale Zimmerman (1978) described a call on a December Myiarchus in New Mexico that he transcribed as reeeeeee. He wrote it "was not a truly clear whistle throughout, but possessed a slightly rough quality at the beginning of the call." Zimmerman also wrote that the call "was a rather thin, almost whistled, rising single note — not emphatic (like the calls of Great Crested M. crinitus) yet not plaintive in quality (as in that of Olivaceous Flycatcher [this bird now called Dusky-capped]). It was a little longer than the ordinary "wheep" note of M. crinitus and was vaguely reminiscent of that. It was, however, much softer, more nearly whistled, and with no hint of a terminal consonant sound;" (emphasis in the original). I think it likely that Zimmerman's bird was a Nutting's, as was published it probably was (he was unable to obtain confirming visual features of importance). Will Russell, in Internet commentary on the Arizona bird, said the call was "a sharp weenk like a muted Great Crested and far from the "police whistle" or soft pwhit notes from Ash-throated. He also said that "the call was uttered rarely (a seasonal silence?)." I find it interested that those hearing presumed Nutting's state the "classic" call is only "vaguely" like a Great Crested, or "like a muted Great Crested," rather than the conventional wisdom of a strong similarity.

In the following chart, I've tried to condense what this literature has to say about vocalizations, lining up similar ones in the same row. All authors recognize that the birds have "diagnostic" calls and other non diagnostic calls that are similar to other species (but apparently the birds can tell even these apart). Both sexes give the four basic calls; there are combination of calls in Ash-throated and Nutting's that extend the number of vocalizations to 6 or 7 (Lanyon 1961, Cardiff & Dittmann 2002). There are also male only dawn songs on the breeding grounds that I do not consider here.

Although I did some comparing of spectrograms to come up with this summarized chart, it seems likely some points were misinterpreted. Lanyon (1961) did not use words to describe most of his calls, so I tried to find comparative words out of Howell & Webb (1994, 1995) but could have assigned them wrongly. So it is the Nutting's column that is most speculative here.

In addition, Joe Morlan has posted sound files from Chris Benesh and Alvaro Jaramillo that include Nutting's Flycatchers in Mexico. I am personally unable to access those files (no broadband at home, no sound card at work) but those who can do so have found them useful.

Words are a difficult tool to use to describe vocalizations, and most everyone will use different words for the same calls. For example, five of us listening to the Santa Cruz bird on 3 January described the calls this way:

• "a soft pwip to a sharper pweep (essentially variations on the same call)" [Steve Gerow]
• "low pitched, variously quiet to loud, single note, pweet, bweep, pweep, or weep, not much of the wh sound as implied by others as wheep" [David Suddjian]
• "poet poet; short but not very loud. This word choice .. is misleading, as I did not hear a two-syllabled note. Later (when Roberson showed up), the call was wick wick wick - not with up-slurred wheet. No long ee sound." [Dan Singer]
• distinctive sharp, short, pwik call, given often when it was vocalizing, sometimes could be rendered as wick!  This call ... was closer to a cross between Downy Woodpecker and Least Flycatcher (i.e., between a pik and a whit)."¹ Also often doubled or trebled pwik-pwik-pwik. [Don Roberson]
• a sharp "pweeep" ....  recalling a quiet Brown-crested Flycatcher;" also a "series of abrupt rising notes" [Joe Morlan]

• "both Joe's description .. as a sharp qwip and Don's described of it as a distinctive sharp, short, pwik ... sound perfectly like the call that I heard repeatedly (personally described as chirk, drrt, or possibly wip). In fact, thinking back about it.... I may even go with a sharp, whistled wip! as the best description for the call I heard."

Steve Gerow and David Suddjian spent more time with this bird than anyone else, and they heard two or three other calls, described as follows

• "when it first started vocalizing in the morning (before it came into view), I once heard a rolling call among the pweeps which seemed to fit the description of this sort [of rolling call] in Lanyon (1961)" [Steve Gerow]
• "while is was perched above us, it once gave a much louder, sharper, and longer variation of its normal call, an upslurred pwheeet, that may be what some describe as the 'typical' call" [Steve Gerow]
• "several calls that were louder and sharper than what I've mostly been hearing from it (which were weep or pweep type calls). The louder calls could be characterized as a variation of the calls I'd heard before. I noted them as pweeet! or pweeek! This call was more drawn out than the other call and a bit upslurred." [David Suddjian]
• "once as the bird flew in from some distance, [it gave a] quick call something like pit-tur-brit, with emphasis on the last syllable and with a rolling quality reminiscent of the calls of Ash-throated Flycatcher" but "I don't think I would characterize that one-time call as rough or burry...but it is hard to get too exact about a quick call heard once." [David Suddjian]

BEHAVIOR: One of the immediately curious aspects of the Santa Cruz bird was that it spent most of its time eating olives. It would take them off the olive tree, or, just as frequently, pick them off the ground under the tree. It would "shake it and beat it against a branch or the ground, in a manner of a large flycatcher or tanager dealing with an insect" (to quote David Suddjian), and then swallow them whole. The pit was later regurgitated. "It dropped a number of olives in the process, either by choice or unintentionally. It ate olives that were ripe and firm, as well as those that were over-ripe and partly shriveled;" [D.L. Suddjian]. For some time, it appeared that this was entirely "an olivacecous flycatcher."²

I thought it very odd that a flycatcher in the genus Myiarchus should be fruit-eating (although I was aware that flycatchers of other tropical genera ate fruit regularly). David Suddjian, however, who has spent hours with the bird, also "saw it forage for insects or other invertebrates several times." He estimated the foraging time was split 70% of olives and 30% of insects. He eventually found it also eating berries at a cottoneaster tree.

Both the Nutting's in Arizona and the Nutting's in southern California spent a lot of time eating fruit. However, all 3 North American species of Myiarchus are known to ingest some fruit on their wintering grounds (Lanyon 1997, Cardiff & Dittmann 2000, 2002). While Ash-throated is almost exclusively a carnivore on arthropods in the breeding season, it will eat fruit in migration or the wintering grounds. Vegetable matter was in 13% of 91 stomachs collected in California between April-December (the percentage in winter not quantified; Cardiff & Dittmann 2002).

The other pages in this project are:

Footnotes

¹ Later I learned that some "helpfully" retranslated this reference of a cross between a Downy Woodpecker and a Least Flycatcher to conclude the bird gave a standard Ash-throated whit or wit. Actually, we were listening to a Downy calling in the background -- a sharp pik -- and the sharpness of the flycatcher call was in that category, but there was a 'pw' sound, so that's where a throw-away reference to Least Flycatcher came in. The call was much closer to the woodpecker. It had no 'wetness' to it like a call rendered whit or wit would have.

² Okay, this is a pun. If you don't get it, you are too young. I don't know if puns are worthy of "credit," but this is a David Suddjian pun. [If you are still scratching your head, please recall that Dusky-capped Flycatcher was known as Olivaceous Flycatcher some decades ago (for its color, not diet), and this Santa Cruz bird was first thought to be an Olivaceous, err, Dusky-capped Flycatcher.

• Bowers, R.K., Jr., and J.B. Dunning, Jr. 1987. Nutting's Flycatcher (Myiarchus nuttingi) from Arizona. Amer. Birds 41:5-10.
• Cardiff, S.W., and D.L. Dittmann. 2000. Brown-crested Flycatcher (Myiarchus tyrannulus) in The Birds of North America, No. 496 (A. Poole and F. Gill, eds.). The Birds of North America, Inc., Philadelphia, PA.
• Cardiff, S.W., and D.L. Dittmann. 2002. Ash-throated Flycatcher (Myiarchus cinerascens) in The Birds of North America, No. 664 (A. Poole and F. Gill, eds.). The Birds of North America, Inc., Philadelphia, PA.
• Devillers, P. 1971. The alleged occurrence of Nutting's Flycatcher in Baja California. Calif. Birds 2:140.
• Dickerman, R.W., and A.R. Phillips. 1953. First United States record of Myiarchus nuttingi. Condor 55:101-102.
• Dittmann, D.L., and S.W. Cardiff. 2000. Let's take another look: Myiarchus flycatchers. LOS News 193:3-10. Louisiana Ornithol. Society, Baton Rouge.
• Howell, S.N.G., and S. Webb. 1994. Field identification of Myiarchus flycatchers in Mexico. Cotinga 2:20-25.
• Howell, S.N.G., and S. Webb. 1995. A Guide to the Birds of Mexico and northern Central America. Oxford Univ. Press, New York.
• Lanyon, W.E. 1961. Specific limits and distribution of Ash-throated and Nutting flycatchers. Condor 63:421-449.
• Lanyon, W.E. 1997. Great Crested Flycatcher (Myiarchus crinitus) in The Birds of North America, No. 300 (A. Poole and F. Gill, eds.). The Birds of North America, Inc., Philadelphia, PA.
• Murphy, W.L. 1982. The Ash-throated Flycatcher in the East: an overview. Amer. Birds 36:241-247.
• National Geographic Society. 1999. Field Guide to the Birds of North America, 3rd ed. Nat. Geogr. Soc., Washington, D.C.
• Phillips, A.R., and W.E. Lanyon. 1970. Additional notes on the flycatchers of eastern North America. Bird-Banding 41:190-197.
• Pyle, P. 1997. Identification Guide to North American Birds, Part I: Columbidae to Ploceidae. Slate Creek Press, Bolinas, CA.
• Zimmerman, D.A. 1978. A probable Nutting's Flycatcher in southwestern New Mexico. West. Birds 9:135-136.

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