Digging Deeper into Tanner: Part 1 of 3 – Tree SizePosted: March 17, 2016
Although it is thematically quite different, this series of posts is rooted in my recent reexamination of my feeding sign hypothesis that culminates here. It was also inspired by my recent and much closer look at Tanner and the Singer Tract, new insights gleaned from old material, and the input of others that shaped the previous post. My original plan was to make this entry the last in the previous series, but since it has grown to over 5,000 words and addresses different issues, I decided to break it in three and will post the next two installments soon.
I’ve been engaged in an extended dialog with a biologist who is familiar with all the Ivory-billed Woodpecker literature and knows Tanner’s writings specifically. Our back and forth is the primary reason for the long interval between the previous post and this one. This person provided some very important insights that will be included in these posts. At the same time, we have a few points of disagreement. In the interests of transparency and allowing readers to draw their own conclusions, these points of disagreement will be disclosed in the text.
Ivory-billed Woodpecker foraging behavior and diet and what separates this species from Pileated Woodpecker and other North American woodpeckers are issues that have been hotly contested for years. In my view, ivorybills could (and presumably do) forage on any species of tree in any decay condition. However, Campephilus anatomy is specialized, and the only quantitative, observational data that exist on what this species does while feeding young (Tanner 1942) suggest some specialization was in fact occurring at least at the Singer Tract from 1937 to 1939. The problem is that many of the prey items (specifically identified in Tanner and emphasized by others since Tanner’s study), even during breeding, do not seem to match up well with the foraging substrates documented by Tanner as most used by ivory-bills feeding young.
In my view, there are some discrepancies between what Tanner observed and reported and the physical evidence he collected during his study related to ivorybill feeding. I also think there may be discrepancies between Tanner’s observations and those of others from the Singer Tract. At least one thing is clear, Tanner’s observations and the photographic record differ markedly from some of his later recollections. In addition, the monograph itself is sometimes ambiguous, as is evidenced by the disagreements mentioned above. It should become clear that the ambiguity and occasional lack of clarity in Tanner’s monograph have led many, myself and my collaborator included, into misinterpretations. We hope that this series of posts will shed more light and clear up some of the ambiguities.
As most readers already know, Tanner’s observations were restricted to one (and the same) family group each of the three breeding seasons during his study. While a sample size of essentially one family group would normally be a serious constraint for comparing with other information, it is important to point out this information represents the only detailed information we have on prey, foraging behavior, and breeding success for ivory-bills, keeping in mind this family successfully fledged young each of these three years. So the data and information Tanner reported on is directly relevant for understanding what was important for successfully fledging young under the conditions found at the Singer Tract during the late 1930s, but as Tanner himself pointed out in his monograph “…the conclusions drawn from them will not necessarily apply to the species as it once was nor to individuals living in other areas.”
Regarding the observations of others on the Singer Tract, I’ll begin with what may have been the last sighting of the John’s Bayou male. In August 1941, George Bick saw three ivorybills feeding in an ash flat near Sharkey Road, quite likely between the bridges over John’s and Methiglum Bayous, south and west of the John’s Bayou home range as delineated by Tanner. This is the only area along Sharkey Road that Tanner listed as “Ash Flat” on his 1941 map.
According to Bick, “I immediately stopped the car and noticed two Ivory-billed Woodpeckers perched in two small ash trees about eight inches in diameter, having recently killed tops. Only one of the birds was carefully observed. A bright, white bill, flaming red crest, and large white wing patch were all clearly noted as the bird remained at the tree. The second bird in a similar ash tree was observed less carefully . . . [A third bird] flew from a dying water-oak tree ten inches in diameter which had only a few curled brown leaves. A stripped spot about six by eight inches and about seventy feet from the ground was present on the trunk of this tree. This is thought to be a spot where the birds had been feeding and to represent the characteristic Ivory-bill ‘sign.’ In the immediate area were many ash trees with dead tops. Much of the bark was stripped in patches of varying size. This may possibly be old Ivory-bill feeding grounds.”
Logging had taken a significant toll in the Singer Tract by the time of Bick’s sighting. It’s thus possible that the birds were foraging in a suboptimal area due to logging pressure. Nonetheless, it’s still worth pointing out that Bick’s observations were in habitat and on tree species where Tanner observed virtually no foraging activity during his study (which ended two years prior, in 1939; he had no feeding observations on water oaks and only one on an ash). It’s also worth pointing out that Bick made specific reference to sweet gums (what he called “red gums”) as being abundant elsewhere but absent from this location.
My collaborator suggested that Bick’s inference that this ash flat was an “old Ivory-bill feeding ground[s]” is questionable. He suggested that changes in hydrology due to logging may have led to an ash die-off. He also noted that this was Bick’s only observation during his six month stay in the Tract, indicating that he was either not looking hard for ivorybills and/or that ivorybills were not using the ash flat on a regular basis. He added another caveat: it is important to remember that Bick’s observation was in August, well after the breeding season when even Tanner assumed foraging behavior for Ivory-billed Woodpecker likely expanded to different habitats and tree species than used during the time they were feeding young at John’s Bayou.
It’s interesting to note that the last known roost, where Don Eckelberry and young Billy and Bobby Fought famously said goodnight to a lone female ivorybill in April 1944, was apparently located in the ash flat where Bick saw his birds (W. Barrow pers. comm.). Just a few months earlier, in December-January 1943-’44, Richard Pough found a lone female roosting in the heart of the John’s Bayou range, about a mile north and east. According to Pough, who was convinced she was the last ivorybill in the Tract, this bird only crossed the Bayou once “for a brief visit to some trees a few hundred feet west of it . . . confining its activities to an area of hardly more than one quarter of a square mile, within which there were an unusually large number of dying trees.”
In our most recent conversation, my contributor and I touched on the question of whether Bick’s birds (and presumably the one seen by Eckelberry and the Foughts) were from the John’s Bayou family group. Either way, it’s a potentially interesting wrinkle. If the birds did come from John’s Bayou, this points to a heavier use of the ash flat for a period of years than is suggested by the limited information about the family group after 1939. All other observations – Pough, Peterson, Tanner, and Baker – were in the heart of the John’s Bayou home range, and at least one of those birds was reliably present there until shortly before Eckelberry and the Foughts said goodnight. On the other hand, if Bick’s birds were a different family group, it suggests that more ivorybills were in the Singer Tract in 1941 than is commonly assumed. (It’s worth repeating that Peterson wrote that one ivorybill was seen in December 1946, and the last letter to Tanner directly related to the Singer Tract birds says that game warden Gus Willett saw a pair in November 1948 and mentions other reports from around that time.)
To return to the Bick report: all of the trees seem to be in the smallest of Tanner’s size classes, 3-12″ in diameter. This class comprised 75.1% of the forest but was the source of only 12.7% of Tanner’s feeding observations. Tanner believed that ivorybills prefer larger trees because they “have more dead and dying wood” but his own data on this are ambiguous, and what he characterized as large seems problematic. The assumption about older trees having more dead and dying wood may have been true around John’s Bayou during Tanner’s study, but this is by no means always the case – the pine forests of Florida, for example, where Allen and Kellogg found abundant feeding sign on young dead pines, which are more vulnerable to fire than mature trees. And as pointed out in the previous post, even in the Singer Tract, the Mack’s Bayou home range was mostly second growth, so forest composition there must have been quite different.
There are a couple of ways to interpret this data. It’s true that 87% of the feeding was “on trees that are over a foot in diameter”, but this is somewhat misleading. 13-24″ diameter trees are the second smallest size class. They hardly qualify as large and approaching senescence, yet they account for 49% of Tanner’s feeding observations. It’s also true that, relative to abundance, the Singer Tract ivorybills showed a strong preference for trees in the 25-36″ class, but the abundance/observation ratios for 13-24″ trees and over 36″ trees are nearly equal, with a slight preference for the smaller size class not the largest. Thus, I think it’s equally accurate to characterize the data as showing that over 60% of observed ivorybill foraging was on smaller trees, under 24″ diameter at breast height and to reiterate that the most often used feeding trees were in the second largest size category, not the largest. (Tanner pp. 43-45).
On the other hand, there’s a good argument that the data show Ivory-billed Woodpeckers foraged on trees in the 13-24” class at 2.6 times the availability, in the 25-36” class at 6.7 times the availability, and in the 36” plus class 2.57 times the availability; there were very few trees in this size class, most of them sweet gums and a few Nuttall’s oaks (Tanner pp. 43-45). Contrast this with the 3-12” class, when the trees were 5.9 times more available than used.
A few additional points should be added to the mix. The numbers discussed above are aggregates, and size preferences were not at all evenly distributed among tree species. Fully 20% of Tanner’s total observations involved sweet gums in the 13-24” class, the most fed upon type. On sweet gums, frequency and abundance ratios are similar for the 13-24” and 25”-36” classes (the latter is the second most fed upon type, comprising around 18% of Tanner’s total observations). For Nuttall’s oak, 13-24” and 25-36” trees were approximately equal in abundance, but Tanner observed considerably more frequent feeding on the larger class.
My collaborator argues that it is more important is to recognize that when combining the data on sweet gums and Nuttall’s oaks, they collectively comprised 31.4% of the total forest and 79.3% of the foraging observations. Trees within the 25-36” class made up 31% and trees within the 13-24” class made up 29% of all foraging observations. Almost all of the trees in the 25”-36” class (5.2%) were in fact sweet gum or Nuttall’s oak, but for trees in the 13-24” class (18.3%) only about 5% (or about a third) were of these two species. This further highlights what Tanner described as heaviest use on sweet gum and Nuttall’s oak for the John’s Bayou family group over all other available trees, and a disproportionately high use of the second largest size class relative to abundance. However, this documented use pattern was not to the total exclusion of other tree species or even the smallest size class available.
This last was a point of contention. I took issue with aggregating sweet gums and Nuttall’s oaks, since they grow and mature at different rates. In addition, I think it’s important to highlight the fact that 13-24″ sweet gums were the single most fed upon type both in terms of frequency of observations and ratio of observations to abundance (albeit it by a small margin). As I see it, this undercuts the misinterpretation of Tanner that ivorybills are ‘large tree specialists’, a misinterpretation I think Tanner invited when he wrote, “The reason for Ivory-bills feeding on the bigger trees is that large, old trees have more dead and dying wood. Young trees grow rapidly and are resistant to the attacks of insects and disease.” As trees ‘mature’ their growth slows and becomes less vigorous, decay begins, insects attack them, and woodpeckers come after the insects.” (p. 43).
In light of this misconception, I also think it’s important to reiterate that in the aggregate, the over 36″ size did not show anything near the disproportionately high use of the 25-36″class. In fact, the rate was very slightly higher on the 13-24″ trees.
Regardless of how one interprets this very limited data set, the idea that Ivory-billed Woodpeckers required ‘large trees’ for foraging has become a truism. The reality is considerably more complex.
The next installment will focus primarily on decay class, and the final one will look at prey species. Stay tuned.