Bits and Pieces Part 6: Rethinking Range and Habitat – Implications

If you’ve been following this series (Part 1, Part 2, Part 3, Part 4, Part 5) you know it has focused primarily on preconceptions about ivorybill range and habitat types and how the actual record paints a very different picture from what many of us think we know about the Ivory-billed Woodpecker. As I noted in the most recent installment, if our knowledge of the Ivory-billed Woodpecker were based on the archaeological record alone, we’d think of it as an upland species. Further, we might very well assume that it ranged from the hills of Georgia, to the Alleghenies in Virginia, to central Ohio and west-central Illinois.*

While it may border on heretical to say so, I think there’s a plausible argument that the ivorybill’s range prior to around 1800 extended as far north as the mid-Atlantic states (New Jersey and Pennsylvania on the Eastern Seaboard) and as far north as central Ohio west of the Appalachians. I’m inclined to think this is likely based on a number of accounts including: Peter Kalm (a student of Linnaeus who reported the species was present in New Jersey and Pennsylvania in the 18th-century), Jefferson (1780s) and Nuttall (1840s) who included Virginia in the range, and Gerard Hopkins a Quaker from Maryland traveling to Indiana to meet with the Miami and Potowatami Nations. Hopkins described a female ivorybill at Piqua, Ohio (north of Dayton, elevation 873′) in 1804 (Leese, 2010.)

In addition, there’s the specimen that Wilson reportedly collected near Winchester, Virginia ca. 1810 (Jackson) and the central Kentucky specimen reportedly collected in the 1780s (Jackson, accepted by Tanner in 1989).

November 2021: Edited to add the Wilson record for Winchester, Virginia is apparently erroneous.

As I see it, the tendency to treat these records as suspect is based, at least in part, on post-Civil War or post-Audubon “knowledge” about the ivorybill and its habitat, rather than anything intrinsically implausible about the claims themselves.

At minimum, one of the Ohio archaeological finds dates to the 15th or 16th century, so there’s strong reason to think that the ivorybill’s range extended that far north at the time of contact. North American Native populations began to decline after Columbus’s arrival, and De Soto’s expedition, 1539-1542,  led to the collapse of the Mississippian culture. (De Soto also introduced the hogs that plague the southern forests to this day.) As a consequence, countless acres of formerly agricultural lands throughout the eastern United States were reforested and remained so into the 18th and 19th centuries. There’s little reason to think that the ivorybill’s range would have contracted at a time when the total acreage of potential habitat was increasing.

I’m reminded that tree girdling may have been an important factor. The only counterargument to the foregoing suggestion about the increase in total acreage after De Soto is that Native American agricultural activity declined drastically during that period, so that while habitat acreage increased, habitat quality may not have. Tree girdling and intentional burning likely played an important role in creating good conditions for ivorybills and could conceivably have led to range expansion during the Mississippian period and again temporarily during the first couple of hundred years of European settlement.

Ivory-billed Woodpecker use of girdled trees was noted by several early observers – notably Audubon, Gosse, and Scott (in Florida, later). While researching this aspect, I came across an interesting account from 1840s Central Louisiana, apparently just south of Alexandria (the citizens of that city are described as “chiefly gamblers or cunning speculators, a nest of incarnate devils, who live by cheating the latest comers, and, whenever possible, each other.”) I’m not aware of this account having found its way into ivorybill literature:

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From Echoes from the Backwoods; or, Scenes of Transatlantic LifeCaptain R.G.A. Levinge (1849).

With this as background, I’d like to propose an alternative explanation (or more accurately an alternative group of explanations) for the ivorybil’s decline. If you think, as I do, that the ivorybill has persisted, this may help explain how the species survived and may even provide some hope for its future, even in this era of mass, anthropogenic extinction.

When it comes to the decline and possible extinction, there has been a tendency to look for one or two causes. The IUCN Species Account gives the following reasons:

Logging and clearance for agriculture are responsible for the dramatic decline in numbers and range. These factors are likely to threaten any remaining population. Hunting has also been implicated in the rapid population decline, and it has been proposed that this was the primary cause of its decline, with habitat destruction playing a secondary role, but this theory is contentious (Snyder 2007, Hill 2008, M. Lammertink in litt. 2012).

Tanner emphasized the importance of logging during the post-Civil War era, although several of his data points seem to suggest that ivorybills were disappearing prior to the most active logging dates. He also stated that the ivorybill’s disappearance “coincided at least roughly with a time of active or rapidly increasing logging.” Elsewhere in the monograph, he focused on food supply, and I suspect that this, rather than logging per se was a more important factor in the ivorybill’s decline.

That’s not to say logging was unimportant; it clearly played a major role. To expand briefly on the point Bill Pulliam raised: by the late 19th century, the more adaptable Pileated Woodpecker, had been extirpated in many parts of its range, and many expected it to ‘go the way of the ivorybill’. That didn’t happen, and PIWOs returned to or became more common in many areas (my own included) as farming gave way to suburban development and forested acreage increased as a result. I’d suggest that for the ivorybill, habitat degradation, rather than habitat loss, was what initiated the decline, with extensive logging and then hunting accelerating an already existing trend.

That is to say, a number of additional anthropogenic factors likely played a role in the ivorybill’s decline and dwindling range, especially outside of Florida, where hunting and collecting likely had much greater impacts than elsewhere. Hasbrouck, writing in the 1890s, contrasted the lack of collecting in Louisiana, Arkansas, Missouri, and Tennessee with what was transpiring in Florida at the time. And it’s important to remember that Florida, which retained ‘frontier’ characteristics far longer than other parts of the eastern United States, was ground zero for the killing and collecting of birds – for commercial and ostensibly ornithological purposes. Ivorybills appear to have been more common in Florida than elsewhere by the second half of the 19th century, but it also seems probable that they were far more heavily persecuted there than anywhere else.

I’m hypothesizing that the shrinking distribution was correlated with settlement patterns in the northeastern part of that range and that by the middle of the 19th-century, east of the Mississippi, it had dwindled to the now familiar outlines, such as those shown on the IUCN range map. Screen Shot 2017-12-22 at 3.42.31 PM

The situation west of the Mississippi is somewhat more ambiguous. A specimen was collected at Forest Park, Missouri (near Saint Louis) in 1886, and there are records from west of the map in Texas dating to the early 20th century. Nevertheless, the general trend toward a shrinking range, which was frequently described in the 19th century literature, is clear.

European settlement brought about numerous changes in the land even before wholesale clearing of forests began.

As mentioned briefly in the discussion of tree girdling, Native Americans used fire for agricultural and wildlife management purposes, something that was likely beneficial for ivorybills. As Native Americans were exterminated, pushed out of their original homelands, or confined to small reservations, and as European settlers tried to control or eliminate fires, a significant factor contributing to tree mortality was likely reduced, dramatically.

Fulton’s invention of a commercially viable steamboat in 1807 revolutionized commerce, drastically accelerating the clearing of log jams from many watersheds in eastern North America. It’s fair to say that “widespread removal of instream wood for steamboat routes, timber rafts, and flood control was equally significant in decreasing floodplain sedimentation and river complexity, and in causing a fundamental, extensive, and intensive change in forested river corridors throughout the United States.” (Wohl, 2014.) As with changes in fire regimes, this clearing of log jams likely led to a decline in the number of stressed and dying trees along the riparian corridors that seem to have been so important for the ivorybill.

Perhaps equally if not more important in my view is the extirpation of the beaver. It is almost impossible to overstate the role of the beaver in shaping ecosystems throughout North America, a subject that’s addressed in engaging detail in Frances Backhouse’s Once They Were Hats. Beavers help create conditions that are good for woodpeckers by stressing and killing trees, through foraging and by changing hydrology. I’ve never tried to quantify it, but many, perhaps most, medium to large sized sweet gums in our search area show signs of beaver damage, and many others have been killed or severely weakened by beaver-caused flooding.

While beavers are not native to peninsular Florida, the ivorybill’s dwindling range elsewhere roughly tracks their decline; with extirpation starting in the northeast, moving West, and then South. (Southern beaver pelts were less valuable.) By 1900, beavers had disappeared from most of the southeastern US, and in Tanner’s day, a very small population persisted in the Florida Parishes of eastern Louisiana. Reintroductions began in the 1950s, and beavers are now considered a pest animal in Louisiana. It’s worth pointing out that the introduced beaver population in Tierra del Fuego appears to be benefitting the native Magellanic Woodpecker (Soto et al. 2012).

The resurgence of the beaver throughout the southeastern US is almost certainly producing substantially improved habitat conditions in many places. While the old growth forests may be virtually gone, it’s not inconceivable that ivorybill food sources are considerably more abundant now than they were in Tanner’s day, and if the species survived, conditions may actually be more favorable than they were in the 1930s and ’40s. It’s also worth pointing out that the southeastern United States is one of the few places in the world where forest cover has increased substantially in the 21st century.

It should be clear to readers of this series that the Ivory-billed Woodpecker inhabited a larger range and was able to exist in more varied habitats than most publications on the species suggest. This has implications for searchers and for what is deemed to be suitable habitat. For example, the trail cam images from the old Project Coyote search area were obtained near the edge of a bean field, and the putative ivorybill roost holes were in willows (more on that in my next post). Since ivorybills in the western part of their range seem to have lived in willow and cottonwood dominated riparian corridors, fast growing, short-lived willows might have played an important role in the species’ survival in other areas too, although willow-dominated habitat would be dismissed as unsuitable under conventional standards of habitat appropriateness.

It seems to me that even a slightly higher degree of adaptability would increase both the chances of survival and the likelihood that surviving populations might be overlooked due to preconceptions about habitat “suitability” ; this was doubtless one of the factors that led officials to dismiss the landowner in our old search area. Now that beavers are again abundant in the southeast, habitat that might otherwise have been deemed “unsuitable” may now be able to support ivorybills, even if the forest itself is not very old. While I don’t envision a recovery along the lines of what’s happened to the Pileated since my youth (when seeing my first one was a thrill as much for its scarcity as its beauty), I think it’s possible that ivorybill numbers have been increasing gradually and modestly over the past few decades. There was, of course, fairly intensive searching from around 2000-2010 (though it’s mostly over now), but it may be that the more numerous sightings from this period and afterwards are due to more than just the increased effort.

*The remains found in Native American middens were unlikely to have been trade goods; ivorybill parts seem to have been a valuable commodity for ceremonial use west of the Mississippi but not east of it, and in several cases, the remains found were tarsometatarsi, which would be consistent with use as food:

There is strong physical evidence of  ritual value for woodpecker scalps and bills from the upper Midwest and Plains . . .  Remains of the Ivory-billed Woodpecker can be found in sacred bundles, on pipe stems, on amulets, and with burials among the Native Americans of the region. The evidence comes from the western Great Lakes and the Plains; no
evidence of a particular use of Ivory-billed Woodpeckers has yet been un-
covered from the eastern area of the Great Lakes (Ohio, Indiana, and Michigan).

(Leese, 2006.) Leese also points out (in several of his publications) that there’s no evidence that ivorybill parts other than scalps and bills had any trade value.

A number of these midden records were accepted by Tanner in his unpublished 1989 update.


Bits and Pieces Part 5: More on Range and Habitat Variety

I intended this long-delayed installment in the “Bits and Pieces” series to be the year’s final post. I also intended it to the be final installment in the series. While this may end up as the last post for 2017, I’ve decided to cut this short and save some discussion for what I anticipate will REALLY be the final installment.

Follow the links for Parts 1, 2, 3, and 4.

I’ll begin by reiterating that my focus here is not just on records from outside what’s commonly believed to be the historic range but also on records from habitats that depart from the stereotype that has emerged since the 1940s. Those who have studied the ivorybill’s natural history in depth understand that the species was not limited to Singer Tract-like habitats, and Tanner himself made this clear. Nevertheless, there’s a widespread tendency to think about habitat quality with a somewhat romanticized image of the Singer Tract – a vast contiguous area of old growth or virgin forest with huge trees and a high canopy – as the touchstone.

Ivorybill anatomy is specialized and particularly well-adapted to scaling bark, but the specialization pertains to securing food sources not easily retrieved by other woodpeckers (including Pileated) during the breeding season; it is not a specialization tied to particular tree species or habitat types.

Before moving on to a discussion of other records that don’t fit the Singer Tract model, I wanted to address something Cyberthrush said in an October post linking to this series. He wrote:

One of my hopes for the widespread USFWS/Cornell search was that it would at least narrow any possible IBWO persistence down to a very few (perhaps 2-3) localized areas; instead the failed endeavor left open the possibility of 2 dozen or more (sometimes little-birded) areas that scarce IBWOs might conceivably utilize. The lack of a single Ivory-billed Woodpecker appearing on remote, automatic cameras by now at more traditional and well-searched areas remains a pretty devastating obstacle to hope for the species… unless indeed it has found a home in the canopies of less-obvious, lightly human-trafficked woodlands.

I don’t want to hold out too much false(?) hope for this species, but on the other hand I believe most southeast woodland habitat is rarely birded in any regular or significant fashion and the vast majority of individual woodland birds are never systematically recorded — moreover, the ornithological literature is rife with weak, unscientific conclusions/generalizations/assumptions about bird behavior, and perhaps even bird biology. There’s just a lot we don’t know, while pretending we do.

While I don’t share Cyberthrush’s perspective on the lack of remote camera photos (or canopies), he makes very good points about the overall paucity of birding activity in many southeastern forests (I have never encountered other birders in the field) and about the limits of our knowledge.

With regard to his disappointment in the organized search effort, I suspect his hopes were founded in part on similar internalized beliefs; these beliefs were reflected to some extent in the excellent lists of potential habitat compiled by Bob Russell and Bill Pulliam that were so widely referenced a decade or so ago. That’s not to say the listed locations were unworthy of attention (and some of them have yet to receive much); however, there are many other areas that were not included.

In discussing his search efforts in one of these areas, Bill summed it up very well:

Many forest birds have modified their habitat usage in recent decades and centuries. Chimney Swifts are an obvious example; also quite telling is the Vaux’s Swift which has only begun moving out of old growth and into chimneys in the last few decades. But of more direct relevance, consider the following quotes:

When found they are usually in regions of original forest growth, rarely being seen where the woods have been once cut over. […] As this Woodpecker seems not to possess the faculty of adapting itself to the new conditions created by civilization, it is quite possible that it will not long survive the passing of our primeval forests. T. Gilbert Pearson, Birds of America, 1936.

Its presence in a region is more often revealed by the large cavities it excavates in dead stumps and trunks than by actual observation of the bird itself. […] This species is common only in the wilder parts of its range. Frank M. Chapman, Handbook of Birds of Eastern North America, 1939.

Both of these early 20th Century characterizations are of the habitat requirements of the Pileated Woodpecker. In fact, as we all know, over the intervening decades Pileateds managed to adapt to forest fragmentation and second growth quite well, thank you, and are now widespread in habitats well beyond the range of what was described in the 1930s.

There’s also another fundamental, perhaps somewhat tautological factor at work. If in fact Ivorybills were an obligate old-growth bird, then yes, they are now extinct. If they were never able to utilize second growth and fragmented forest, then there is absolutely no reason to be looking for them now anywhere. But even given the historical accounts of their habits they demonstrated a bit more flexibility than this; and given the actual history of Pileateds and various other forest birds since the industrial revolution, it seems brash and unjustified to presume that the habitat utilization of a 21st Century Ivorybill would be the same as that of one in the 19th Century.

We now know that even within the Singer Tract, ivorybills nested successfully at Mack’s Bayou, an area that was predominantly second growth, and I’d suggest that ivorybills showed more than “a bit more flexibility”, when one looks at the historical record.

As Fangsheath at ibwo.net has pointed out both on the forum and in an email, most of the ivorybill localities referenced by Tanner are from within 20 miles of the coast, and “[t]hese coastal forests are quite different from the Singer Tract.” In many of these coastal areas, tree size would have been smaller, the understory would have been thicker, and the canopy would have been lower due to hurricanes and other factors. In addition, the coastal records may have implications for survival in fragmented habitat types.

There are a number of records from Florida and South Carolina involving offshore islands. While most if not all of these offshore islands would have been covered in old growth forest at the time of collection, reaching them would have required crossing expanses of open water or marsh.

In South Carolina, there were multiple reports from barrier islands into the 1880s. A specimen collected in 1879 or 1880 is now lost. Hoxie, writing in 1918, reported that ivorybills were generally “unpersecuated or harmed by man” and that they had opportunistically fed on the barrier islands, following hurricanes, but disappeared when the food supply was exhausted.

My knowledge about Florida and conditions there is limited, but it seems clear that ivorybills lived and bred in a variety of habitats. Florida is also the largest single source of specimens and probably had the largest ivorybill population in the country. What may also be relevant is their apparent use of of mangrove forests, including potentially some 1-2 miles away from the mainland, especially in the Everglades region. As with barrier islands, use of this habitat may have involved crossing some open water between mainland forests (e.g., Big Cypress, associated forested sloughs, and open pine woods, leading to the Gulf of Mexico) and mangrove forests, both along the coastline and in the area referred to as the Ten Thousand Islands, extending south to where Tanner relayed reports during the 1930s from Shark and Lostman’s Rivers within what is now Everglades National Park.

To end this tour of ‘unexpected’ habitats and ‘extralimital’ records, let’s jump north and west by a thousand miles or so and consider records from Arkansas/Missouri and Virginia.

The Arkansas/Missouri records are from George Featherstonhaugh who explored the area in the 1830s and published his account in 1844 as Excursion Through the Slave States from Washington on the Potomac to the Frontier of Mexico; with Sketches of Popular Manners and Geological Notices. Featherstonhaugh reported seeing ivorybills in two different locations – one from a bottomland area above the confluence of the Ouachita and Caddo Rivers, in the vicinity of present day Arkadelphia, AR. This passage is from Volume 2.

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The second location appears in Volume 1 and is more interesting for the purposes of this discussion, since it involves fire damaged upland forest, likely oak-dominated, with hickory and perhaps some shortleaf pine. It’s not clear whether the site is in present day Arkansas or present day Missouri. Either way, it appears to be on the edge of the Ozark Plateau where it meets the Mississippi Alluvial Plain. As an aside, Featherstonhaugh seems to have had an eye for detail and a dry sense of humor:

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Whether the birds were resident in this area or were merely feeding opportunistically in the fire damaged table-land, the habitat involved bears no resemblance to the Spanish Moss festooned, swamp forest stereotype.

Thomas Jefferson included the ivorybill on his list of Virginia birds found in Notes on the State of Virginia (1785). While some authors have suggested that Jefferson was merely following Catesby, it has also been argued that Jefferson’s list was “a product of his personal observations”. If this is true, Jefferson’s observation would not have been from the Great Dismal Swamp (where Pearson thought ivorybills might be present in the early 20th-century), since Jefferson never went south of Norfolk.

Thomas Nuttall, writing in the 1830s, described the ivorybill as being “seldom seen to the north of Virginia and rarely in that state.” In The History of Ornithology in Virginia (2003), Johnston dismisses Nuttall’s assertion but also points to a much earlier and unambiguous record from an upland site, one that seems to have been otherwise overlooked.

The record is from a Native American midden dating to the early Woodland period, ca. 300 CE (AD). The site, Daugherty’s Cave, was used for millennia. It’s in the western part of the state, far from the coastal plain, at an elevation of approximately 2000′. The context suggests that the remains were not trade goods, and Johnston deems it to be “the only known record of this bird from Virginia”.

I’m struck by the fact that if the ivorybill had gone extinct before 1700, and we only had the scant archaeological record to go on, we’d imagine it to be a bird associated with upland forests. On a more serious note and perhaps one more relevant to the discussion that will follow in the next post, I wonder whether the appearance of ivorybill remains in Appalachian middens around 1800 years ago is related to the spread of maize agriculture and tree kills associated with this new farming technology.

So what inferences can be drawn from all this information and what are the implications? I have some thoughts, not all of them had occurred to me before I began  this exploration, and I hope they’ll inspire readers to reflect and come up with their own new insights.

The first and perhaps most mundane observation is that the range maps with which we’re all so familiar, like the one below from the IUCN, reflect the ivorybill’s status after what was likely a long period of decline, especially east of the Mississippi. Unfortunately, these maps reinforce ideas about habitat requirements and tree species associations that are, at best, oversimplifications.

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For contrast, I’ve updated the map I created showing most of the records discussed in this series.

This is not to suggest that reports from outside this “historic” range should be taken seriously today. The map is a fairly accurate reflection of the reality post-Civil War. The more important questions are why the range started to shrink, probably during the 18th-century, and what this more complex analysis of potential habitat types might imply for the species’ survival. That will be the subject of the next and final installment.


Bark: An Exegesis

Introduction:

According to Tanner, scaling bark was the Ivory-billed Woodpecker’s primary foraging strategy during breeding season in Louisiana. Tanner wrote that the ivorybill is “capable of easily scaling away heavy bark that other woodpeckers could not loosen.” (Tanner 1942). All woodpeckers in genus Campephilus have specific anatomical characteristics that enable them to forage in this specialized way (Bock and Miller 1959). Following Tanner, most post-Singer Tract search efforts have looked for feeding sign as an indicator of presence. Because Tanner’s descriptions are somewhat vague and many of the photographs showing feeding sign are poor, these efforts have tended to focus on decay state and bark adhesion without taking bark characteristics and tree species sufficiently into account. I posit that tree species and bark and wood characteristics are key factors that should be considered. I further posit that extensive bark scaling on live and recently dead hickories (genus Carya) may be beyond the physical capacity of the Pileated Woodpecker.

Discussion:

As all regular readers know, I’ve been somewhat obsessively focused on bark and bark scaling since my earliest years of ivorybill searching. The reason for this interest is simple: it’s how Tanner found ivorybills or inferred their presence when he couldn’t find them (Tanner 1942). Unfortunately, as discussed in a number of posts, Tanner’s descriptions are somewhat opaque, and most of the published images of feeding sign, including those in the monograph, are not very illuminating. Indeed, some of them are consistent with pileated work that we’ve documented. Plate 8, shown below, is a prime example. The caption reads, “Ivory-bill feeding sign on a slender limb”.

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Tanner’s Plate 8, Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

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Pileated Woodpecker feeding sign on a slender limb

Early on in my study of this subject, I hypothesized that certain kinds of bark scaling on hardwoods might be beyond the physical capacity of the Pileated Woodpecker. I still believe this to be true, a view that is supported by what we’ve documented for pileated and by numerous examples of pileated scaling from online sources. At the same time, the details of what types of work might belong in this category have shifted somewhat, especially as it has become clear that Pileated scaling can look like what’s shown in Plate 8 and that pileateds will scale bark from recently dead sweet gums.

This is not to suggest that ivorybills never scale small and medium-sized branches in a similar manner. According to Tanner they did so frequently; however, I have been focused on what may be diagnostic for ivorybill. It seems likely that there is considerable overlap between ivorybill and pileated work when smaller branches are involved (at least on sweet gums).

The sequences we obtained showing pileateds scaling a sweet gum limb have inspired me to look more deeply at the characteristics of hardwood bark and pursue some research avenues that I hadn’t considered previously. I’ve linked to some of the sources in recent posts, but I’ve had some additional insights that seem important enough to share. Every time I think I’ve run out of things to say on the subject, something new crops up.

Like virtually everyone else, I’ve followed Tanner and focused on two bark characteristics, “tightness” and thickness, but it recently struck me that other features might be important as well. And the literature, mostly from the lumber industry, supports this idea.

Tanner suspected that the Singer Tract ivorybills preferred sweet gums and Nuttall’s Oaks because the bark is thinner, and the thinner barked limbs had “more borers” than thick barked ones. While abundance of food was likely a factor, I suspect that, at least with respect to sweet gums and possibly Nuttall’s oaks, ease of scaling and access to food played a role.

It’s important to point out that in live trees, hardwood bark adhesion varies seasonally, with bark becoming tighter during dormant stages and looser (with considerably less variation from species to species) during the growing season. Bark is often if not always tighter on recently dead trees than on live ones (Stokland et al. 2012).

In addition, “The structural and chemical traits of dead wood, inherited from the traits of living trees, are also major drivers of wood decomposition and these traits vary greatly among tree species.” (Cornellisen et al. 2012). The authors of the linked paper point out that other factors, including size and site, can also contribute to the way that bark loosens post-mortem, but specific traits seem to be paramount, especially since the scaling we deem to be suggestive, whether on standing or downed wood, is on trees that are alive or are recently dead. Because the scaling has a very distinctive appearance, we also deem as suggestive hickory snags and stubs that appear to have been scaled some years ago, even if they are in a considerably later stage of decay overall. Bark attached to hard wood on these longer dead stubs and snags often remains tight for 3 or more years after death.

A 1978 report, entitled Bark and Wood Properties of Pulpwood Species as Related to Separation and Segregation of Chip/Bar Mixtures examined bark morphology and strength properties in 42 different pulpwood species and identified factors that impede the mechanical removal of bark from logs. These include: cellular structure, bark adhesion, bark strength, bark toughness, wood toughness, specific gravity/density, and moisture content. (Institute of Paper Chemistry 1978) One caveat about this report: a subsequent paper gives the sample size for each species, and in many cases (including sweet gums) it was only 2 (Einspahr et al. 1982)

It may be counterintuitive, but the authors found that shagbark hickory was far and away the most difficult bark to remove. (The tightly adhering layer is thin, beneath the dead bark that gives the species its shaggy appearance.) One key finding was that:

“Morphologically, the presence of fibers increases inner bark strength and, when sclereids (a type of cell) are present, bark strength is decreased. Inner bark strength, in turn, has a major influence on hardwood wood/bark adhesion. The multiple regression equation employing wood toughness and inner bark strength accounts for 72% of the wood/bark adhesion variation encountered.”

Sclereids are virtually absent in hickories (Nanko 1980) and a few other species that don’t approach the hickories in bark strength and bark and wood toughness (Eastern cottonwoods, yellow poplars, white ashes, and black willows). These tables are particularly illustrative:

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woodchart

Shagbark hickories are the extreme outlier in this study, in terms of adhesion, as well as in terms of inner and outer bark toughness and strength; there are very few shagbarks in our search area, and we have never found scaling on one. I have been unable to find specific information about bitternut hickory bark strength or toughness, but the industry’s debarking problem applies to all species in the genus Carya due to the near absence of sclereids in conjunction with the other factors. Moreover, the industry does not differentiate among hickory species (Timber Mart South 2016). This 1996 paper is worth quoting at length in this regard (full text is not readily accessible):

The amount of published literature dealing with hickory debarking is very limited. Often it is only mentioned as an example of one of the hardest tree species to debark. One study quantified this by measuring the strength of the bark-to-wood bond of 42 hardwood species, including hickory. According to Einspahr et al., the dormant season bark-to-wood adhesion for hickory is greater than 3000 kPa, which is a tenfold difference from the growing season and nearly three times as great as the dormant season wood/bark adhesion for quaking aspen (Populous tremuloides, L.), a species considered to be extremely difficult to debark in the northern United States.

Einspahr et al. also microscopically examined the failure zone in an attempt to correlate morphological differences with bark-to-wood adhesion. For hardwoods in general, they found that during the growing season, failure occurred in the cambium or in the xylem just inside the cambial zone. Conversely, dormant-season failure occurred in the inner bark. They also found that fibers in the bark increased the inner bark strength while sclereids decreased inner bark strength. Hickory bark can contain between 15 to 20 percent fiber and contains less than 0.05 percent sclereids.

While these studies have confirmed that hickory is difficult to debark, they have not addressed possible solutions to the problem. As a result, hickory is often left behind during harvesting, reducing the total usable fiber from a given stand and, over time, increasing the percentage of the species in the hardwood resource, compounding the problem of future harvests.

When a tree dies, the bark eventually loosens and detaches naturally as the cambium decays. After felling, the cambium remains alive until it has consumed all available food or dries out. Moisture loss, while causing cambial death, initially greatly increases the strength of bark attachment because additional bonding between fibers occurs as the secondary valence bonds with water are broken (Belli 1996).

Thus, even though hickory bark adheres less tightly than sweet gum bark during the growing season, it seems likely that it’s harder to scale year round, given its much greater wood and bark strength and toughness. It is also clear from my observations that sweet gum bark loosens far more rapidly than hickory bark post mortem. Note that we have found fresh scaling on both live and recently dead hickories.

Based on specific gravity of the bark – averaging 0.72 for shagbark and 0.60 for bitternut – and bark moisture content – averaging 34% of dry weight for shagbark, and 60% for bitternut – it seems likely that bitternuts are somewhat easier to debark than shagbarks but considerably harder to debark than virtually any other tree species in our search area.

Comparing bitternut hickories to other species, most oaks have a considerably higher average moisture content in their bark (Chestnut and Southern red, including Nuttall’s oaks, are exceptions) and similar specific gravities. Sweet gum bark has an average specific gravity of 0.37 and an average moisture content of 91% of oven dry weight. (Schlaegel and Willson 1983, Miles and Smith 2009). But oaks and sweet gums have sclereids, and sweet gums and all tested oak species score far lower on bark toughness and strength than shagbark and, by inference, bitternut hickories. Sweet gums and the tested oak species are fairly similar in these regards, but I suspect that the higher density and lower moisture content in oak bark makes it harder to scale and may mean that oak bark adheres more tightly than sweet gum bark for a longer period after death.

I posit that when it comes to woodpecker scaling, dormant season bark adhesion, inner and outer bark strength, and inner and outer bark toughness are all relevant factors. We know that Pileated Woodpeckers remove sweet gum bark with some difficulty and that even on medium-sized limbs, they are not consistently able to remove bark cleanly down to the sapwood. It’s also clear that bitternut hickory bark is very difficult to remove, second only to shagbark hickory in our search area. This further reinforces my view that the work on hickories is not the work of Pileated Woodpeckers.

Click here and here for examples of the hickories that are scaled in a manner we hypothesize is diagnostic for Ivory-billed Woodpecker. Also be sure to watch this YouTube video of a Crimson-crested Woodpecker (Campephilus melanoleucus) foraging. (Thanks to Phil Vanbergen for finding the clip and the scaled hickory at the second link.) I’m reposting the link to the video here because I think it very clearly illustrates many of the characteristics we associate with Ivory-billed Woodpecker work on hickories, although the species of tree being fed on is unknown. Note the striking similarity in appearance and also that the work of the substantially smaller billed Crimson-crested is not as clean around the edges as the work we’re ascribing to ivorybills.

There were no bitternut hickories in the Singer Tract, but there were congeners – pecans and water hickories. Tanner observed ivorybills scaling on these species twice and digging once. For pileateds, there were 4 instances of digging and none of scaling, as opposed to 5 scaling and 9 digging on sweet gums. The relative abundance of water hickory and pecan at Singer was 2.7%; approximately 10% of the trees in our search area are hickories, and hickories are second only to sweet gums in terms of the number of scaled trees we’ve found. While Tanner’s is obviously a minuscule data set, it may support the hypothesis that live and recently dead Carya bark is too tough for pileateds to scale extensively, if at all.

There are a number of hardwood species found in potential ivorybill habitat that are somewhere between sweet gums and hickories in terms of how easily scaled they may be and how soon after death bark decay and loosening set in – eastern cottonwoods, black willows, water tupelos, some oak species, red maples, green ashes, honey locusts, persimmons, and elms – in these species, it seems likely that close examination of the scaling and bark chips can provide some clues.

Conclusion:

Previous Ivory-billed Woodpecker searches have focused on bark adhesion and state of decay when considering scaling as possible foraging sign. Bark morphology, dormant season adhesion, inner and bark outer strength, and inner and outer bark toughness, and wood toughness are all relevant to the ease with which bark can be scaled from live and recently dead hardwoods. Specific gravity and moisture content are also factors. Bark from trees in the genus Carya is difficult to remove industrially, and members of this genus are likely the most difficult trees to scale throughout the historic range of the ivorybill. Since Pileated Woodpeckers scale sweet gum branches with some difficulty and do not consistently remove bark down to the sapwood, it may be beyond the physical capacity of Pileated Woodpeckers to scale hickories extensively and cleanly, while leaving large pieces of bark behind. Extensive work on hickories that has a distinctive appearance may be diagnostic for ivorybills; this distinctive appearance of this scaling may also be the key to recognizing Ivory-billed Woodpecker foraging sign on other species.

Lagniappe:

This may be no more than an aside, but it may be a relevant data point. I recently observed a Pileated scaling briefly on a live 14″ DBH Norway maple in my yard near New York City. The photos show that the sap is flowing. The appearance of the scaling is exactly what I’d expect for Pileated, with strips about half an inch across. Norway maple may be a decent stand-in for sweet gum; while its bark has a higher specific gravity, 53 as opposed to 37, the moisture content of the bark is almost identical, 91% as opposed to 90%.

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References Cited:

Bock, Walter J. and Waldron Dewitt Miller, The Scansorial Foot of the Woodpeckers, with Comments on the Evolution of Perching and Climbing Feet in Birds, American Museum Novitates, #1931, 1959

Belli, Monique L., Wet storage of hickory pulpwood in the southern United States and its impact on bark removal efficiency, Forest Products Journal. Madison 46.3 (Mar 1996): 75.

Cornelissen, Johannes H.C., Ute Sass-Klaassen, Lourens Poorter, Koert van Geffen, Richard S. P. van Logtestijn,Jurgen van Hal, Leo Goudzwaard, Frank J. Sterck, René K. W. M. Klaassen, Grégoire T. Freschet, Annemieke van der Wal, Henk Eshuis, Juan Zuo, Wietse de Boer, Teun Lamers, Monique Weemstra, Vincent Cretin, Rozan Martin, Jan den Ouden, Matty P. Berg, Rien Aerts, Godefridus M. J. Mohren, and Mariet M. Hefting, Controls on Coarse Wood Decay in Temperate Tree Species: Birth of the LOGLIFE Experiment, Ambio. 2012 Jul; 41(Suppl 3): 231–245.

Einspahr, D.W, R.H VanEperen, M.L. Harder et al. Morphological and bark strength characteristics important to wood/bark adhesion in hardwoodsThe Institute of Paper Chemistry, 1982: 339-348.

Institute of Paper Chemistry, Project 3212, Bark and wood properties of pulpwood species as related to separation and segregation of chip/bark mixtures, Report 11, 1978.

Miles, Patrick D. and W. Brad Smith, Specific Gravity and Other Properties of Wood and Bark for 156 Tree Species Found in North America, United States Department of Agriculture, Forest Service. Northern Research Station, Research Note NRS-38, 2009.

Nanko, Hiroki, Bark Structure of Hardwoods Grown on Southern Pine Sites (Renewable Materials Institute series), Syracuse University Press, 1980.

Schlaegel, Bryce E. S and Regan B. Willson, Nuttall Oak Volume and Weight Tables, United States Department of Agriculture, Forest Service. Southern Research Station, Research Paper SO-l 86, 1983

Siry, Jacek, ed., Species Detail Report, Timber Mart-South, 2016

Stokland, Jogeir N., Juha Siitonen, and Bengt Gunnar Jonsson, Biodiversity in Dead Wood, Cambridge University Press, 2012

Tanner, J.T. The Ivory-billed Woodpecker,National Audubon Society, 1942.

Thanks to Fredrik Bryntesson, Steve Pagans, Chris Carlisle, and Bob Ford for their help with this post.


Feeding Sign: Further Reflections and Clarifications

I’ve created a page that expands on this post and should provide a good introduction to our thinking about the three most intriguing types of feeding sign, with images from the second and third categories. Think of this post as the shorter version.

Over the years, I have written a great deal about bark scaling and the types of work I think are diagnostic for Ivory-billed Woodpecker; however, I don’t feel that I’ve been effective enough at conveying the nuances of what I look for in situ. I’m going to try a somewhat different approach using images that have been posted previously. I will be focusing on the category of bark scaling that I think is most compelling for Ivory-billed Woodpecker. I hope that the tiled mosaic layout will make it easier to get my points across.

The scaling I find most compelling for ivorybill is on hickories, mostly or all bitternut hickories (Carya cordiformis).  This work represents a relatively small subset of the suspected ivorybill feeding sign we’ve found, as would be expected given that under 10% of trees in the area are hickories. It is not the type of foraging sign that Tanner emphasized, and I’m not suggesting that scaling on boles is the ivorybill’s predominant foraging strategy. I emphasize this work because it has a distinctive appearance, one that differs dramatically from presumed Pileated Woodpecker foraging sign on the same species.

The above images show presumed Pileated Woodpecker work on a recently dead bitternut hickory found last February. It seems reasonable to infer that this is Pileated work because of its extensiveness and the abundance of fresh chips at the base of the snag, suggesting the work was recent and was done by a large woodpecker. Some readers might be inclined to think of this as “scaling”, when in fact it is shallow excavation. The small size of the chips and the fact that some of the chips are sapwood, not bark, support this idea. Contrast the roughened appearance of the sapwood with the clean bark removal in the images below. Also contrast the extensiveness; while the work shown above involves fairly large areas, it pales in comparison to the very extensive scaling shown below.

Edited to add: I think squirrels can also be ruled out for this work due to the involvement of the sapwood, apparent bill marks, and the presence of insect tunnels.

I think that all of the images immediately above show Ivory-billed Woodpecker work, most of it fresh. The similarity in general appearance from tree to tree should be self-evident. This type of scaling can be found from within approximately one foot off the ground to the upper parts of boles. Large Cerambycid exit tunnels are visible in the sapwood. Bark chips, when found, were large, and the only hints of excavation involved targeted digging to expand the exit tunnels. It’s worth noting that the Hairy Woodpecker in the trail cam photo above spent several minutes removing a quarter-sized piece of bark. The Pileated Woodpecker also spent several minutes on the tree; it pecked and gleaned and looked around but did no excavating or scaling.

Bitternut hickory wood is “hard and durable” and the bark is hard and “much tighter than on most other hickories.” The bark is sometimes described as being thin, but this appears to apply to young trees only. On mature boles, it can be .5″ thick or more.

Due to these qualities, the decay process for hickory snags is often more gradual than for other species, especially in higher and dryer areas. This means that the wood can stay hard and the bark remain tight for as long as three years; such is the case with the tree shown in the penultimate image above. This has enabled me to do periodic checks on some of the snags, and in most instances, they have shown little or no indication of further woodpecker work for extended periods, until the wood starts to soften and excavation becomes easier. Whatever is doing this work seems to be hitting the trees once or twice without returning or, less frequently, to be making visits several months apart. I think this suggests a thinly distributed, wide-ranging species as the culprit, and in my experience, Pileated Woodpeckers tend to return to feeding trees on a regular basis.

In my view, this very specific type of work is diagnostic for ivorybill and is beyond the physical capacity of the Pileated Woodpecker. I’d suggest that similar appearing work on other tree species should be considered strongly suggestive. When it comes to the high branch work that Tanner emphasized, it is more difficult to rule out Pileated Woodpecker. As discussed in several recent posts, this type of foraging was the predominant one during breeding season and immediately after fledging young, at least for the John’s Bayou family group. Thus, for work higher on trees, where bark is thinner and tightness cannot be assessed, abundance is likely a key indicator. From this perspective, it may be significant that our friends the Carlisles who are searching in the Pascagoula area have found only two sweet gums with work that I consider to be intriguing and consistent with what’s described in the literature. By contrast, I found over 50 such trees in our area in the 2015-2016 season alone. Whether or not there are ivorybills in the Pascagoula, the difference between the Carlisles’ observations and ours is dramatic and suggests that something unusual is going on in the Project Coyote search area.

 


Insights, Ants, and Old Growth: a Nuanced View of the Ivorybill’s Decline and Possible Survival

I’ve just finished reading Tanner’s dissertation and have gained some new insights into topics that have been discussed in a number of earlier posts.

Conventional wisdom, following Tanner, holds that the Ivory-billed Woodpecker’s decline and possible extinction were caused by habitat loss, specifically the logging of old growth forests during the 19th and early 20th centuries. Birdlife International’s fact sheet on the species suggests “that large contiguous tracts of mature woodland would be required to support a viable population”, referencing Jackson 2002. Snyder et al. have proposed an alternative hypothesis that “human depredation was the primary factor.”  (p.9).

Tanner’s model depends on the idea that food supply was the limiting factor on ivorybill populations, because the species is highly specialized, and that old growth conditions were optimal or essential. While Tanner was aware that ivorybills bred successfully in an area that was predominantly second growth, at Mack’s Bayou, he glossed over this fact in the monograph, and became more dogmatic about old growth as a requirement in later years.

Snyder and some others have contended that the ivorybill is a generalist. According to Snyder, “the data available on diet and foraging methods simply do not provide compelling evidence for strong feeding specialization.” Snyder goes on to suggest that “[i]ts apparent skill in exploiting recently dead timber, coupled with its ability to feed in a variety of other ways, may even have given it some significant foraging advantages over the pileated woodpecker, a species apparently much less capable of bark stripping. Indeed, the pileated woodpecker, like other Dryocopus woodpeckers, may well be more of a food specialist than any of the Campephilus woodpeckers.” (p. 37).

As I see it, there are elements of truth in both models, but neither is complete. In addition, I think that each model relies on at least one flawed premise.

The old growth/virgin forest component of Tanner’s model fails to account for the facts that the Singer Tract population was dwindling even before logging began in earnest and that birds appear to have remained in the Tract until well after it had been extensively logged. Tanner suggested another possibility, “perhaps the greatest factor reducing the rate of ivorybill reproduction is the failure of some birds to nest. One reason for their not breeding is immaturity, for it is probable that ivorybills do not nest until they are two years old. Another possibility is that the quantity of food available to the woodpeckers may determine whether they will nest or not.” (p. 83).

Tanner struggled to account for the fact that the ivorybill population at Singer was dwindling by the mid-1930s, even though overall habitat quality had, if anything, improved relative to what it had been a few decades earlier. He attributed the higher relative abundance in previous years to tree mortality due to fires that took place in 1917 and 1924. Tanner also recognized the probable importance of fire in the pre-contact era, although he seems to have been unaware of the ways pre-contact Native Americans used fire, both for agriculture and habitat management. (The impacts of Native American fire use were almost surely different from what occurred in the 20th century Singer Tract).

Neither Tanner (whose study predates the emergence of the discipline) nor Snyder, take environmental history sufficiently into account. There had been major ‘changes in the land’ long before large scale logging began in the southeast and before the reports of local abundance on which Snyder relies. These changes include: the post-contact collapse of Native American civilizations, the introduction of European plant and animal species, the clearing of log jams on major and secondary North American rivers, habitat fragmentation due to the plantation economy, and the near extirpation of the beaver.

All of these elements likely contributed to a major decline in ivorybill populations. Ivory-billed woodpeckers likely concentrated locally in response to major disturbances, regardless of whether forests were old-growth or advanced second-growth, and this type of specialization caused birds to congregate, making it easier for collectors to kill them in large numbers in short periods of time. Snyder likely misinterpreted this collection of large numbers of Ivory-bills in short periods of time as reflecting a greater regional abundance. In contrast, and more consistent with Tanner, this ecological response to disturbed areas led, in some places, to the collectors extirpating regional populations.

In the latter part of the 19th century, hunting probably sped the collapse of the remaining population, but Snyder’s claim that available data on diet and foraging methods do not provide compelling evidence of specialization fails to account for the anatomical and other evidence that suggests otherwise. It also fails to account for the Pileated Woodpecker’s far more extensive range and ability to thrive in a wider variety of habitats, including badly fragmented and degraded ones. I made some of the case for specialization in a series of recent posts, but there’s more to add, especially with regard to ants.

In one of those posts, I hypothesized that the inability to exploit ants as a food resource was a key component, perhaps the primary component, in explaining the decline of the ivorybill. A commenter asked whether there’s evidence to support the idea that ivorybills and other Campephilus woodpeckers don’t feed on ants and also whether there’s evidence to support the idea that Campephilus woodpeckers don’t regurgitate.

Adult Campephilus woodpeckers rarely feed on ants but do not feed them to their young. They make frequent trips to the nest with food items stored in the bill or at the back of the bill. (M. Lammertink, pers. comm.) Dryocopus woodpeckers and those in closely related genera (the “tribe” Malarpicini) feed their young by regurgitating, while other woodpeckers do not. (Manegold and Topfer, 2012). I think the capacity of Pileated Woodpeckers to consume ants in large quantities and to feed them to their young is a significant distinguishing factor and that Tanner was correct in suggesting that food supply was a major limiting factor on Ivory-billed Woodpecker populations.

Ants comprise up to 33% of the world’s terrestrial animal biomass. In Finland, they comprise as much as 10%. In tropical forests, the percentage is much higher, exceeding vertebrate biomass by 400%. Tanner’s comparative analysis of available ivorybill and pileated food did not include ants, so Tanner’s comparative estimate of available insect prey – suggesting that pileateds in the Singer Tract had access to approximately four times what ivorybills did – was in fact extremely low.

Tanner’s dissertation concludes with a discussion of Audubon’s ivorybill dissection, something that was omitted from the monograph. While I had a passing familiarity with the Audubon material, I had not looked at it carefully. Nor had I compared his ivorybill and pileated dissections.

Tanner wrote: “The proventriculus is both muscular and glandular. Audubon’s drawings and text indicate that the proventriculus of a Pileated is much larger in proportion to the stomach than is the case in the Ivory-bill.” Audubon described the ivorybill proventriculus as being only minimally wider than the esophagus. By contrast, the pileated proventriculus as “an immense sac, resembling a crop, 2 1/4 inches in length and 1 and 5 twelfths in width,” or nearly three times as wide as the esophagus.

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Audubon’s drawing of Ivory-billed Woodpecker digestive tract showing slightly widened proventriculus.

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Audubon’s drawing of Pileated Woodpecker digestive tract showing large, sac-like proventriculus.

The proventriculus and stomach of one of Audubon’s specimens contained “a vast mass of ants and other insects”. According to Bent, Beal found one pileated stomach that contained 2,600 ants. (Others contained fewer, 153 and 469, according to Sutton.) Thus, it’s clear that even if ivorybills sometimes ate ants, they lacked the capacity to consume them in large quantities, let alone feed them to their young.

This supports Tanner’s view that specialization was a limiting factor on ivorybill populations. I’ve previously suggested that this might apply only to breeding season, but it seems reasonable to infer that it’s a factor year-round, based on the differences in proventricular structure.

All of that said, I’d argue that this specialization should not necessarily be read to include dependence on large tracts of mature, contiguous forest. The data from the Singer Tract suggest that even under these ‘optimal’ conditions, breeding was limited. And the fact that the Mack’s Bayou birds bred successfully in an area of second growth suggests that birds could thrive under ‘suboptimal’ conditions. The extent to which survival might be possible in fragmented habitat is less clear, but Snyder (citing Jackson) refers to the Mississippi population of six pairs in a 19.2 square mile forest that Tanner missed; the tract is less than 1/6 the area of the Singer Tract and is smaller than many contemporary wildlife management areas.

The tract, known as Allen Gray Estate, was west of Skene, Mississippi in Bolivar County; some or all of it is now part of Dahomey National Wildlife Refuge; the US Fish and Wildlife Service Habitat Management Plan for the refuge (2013) states that the forested portion of the refuge comprises 8100 acres and provides this historical information, “Dahomey NWR is located on the grounds of the old Dahomey Plantation founded in 1833 by F.G. Ellis and named after the homeland of his slaves. Much of the land west of the refuge was probably cleared for cultivation around this time. The land went through several owners and was purchased by Allen Gray in 1936. The portion that became the refuge was known as the “Allen Gray Woods”. This was the only significant portion of the plantation still forested.”  This 8100 acre figure is 25% lower than the figure reported by Jackson and Snyder.

While I have been unable to find a detailed logging history of Bolivar County, it is in the heart of the Mississippi Delta, which was known for its plantations. Between 1900 and 1940, Bolivar County was more densely populated than Madison Parish: 39.1 people per square mile as opposed to 18.9 in Madison Parish in 1900, 78.92 as opposed to 22.78 in 1930, and 74.57 as opposed to 28.33 in 1940. Based on population density and the number of towns, it seems self-evident that the habitat in Bolivar County was considerably more fragmented than was the Singer Tract.

Thus, there is good reason to question Tanner’s old growth model as well as the idea that large contiguous tracts of mature forest are required. Similarly, there’s good reason to question Snyder’s argument that hunting rather than specialization was the primary cause of the ivorybill’s collapse.

Efforts to reintroduce the beaver in the southeast began in the 1930s, and the population has been growing ever since. Beavers injure trees by partially or fully girdling them and by altering hydrology, which weakens or kills trees at the edges of the ponds they create. Beaver damage renders trees more vulnerable to infestation by ivorybill prey species, something we’ve observed repeatedly in our search area. In Tanner’s day and in the late 19th century, the beaver was barely a part of the southeastern ecosystem, but by the 1950s, beavers again were playing a role in altering southern forests, whether mature or successional.

If the ivorybill was able to survive the logging of the last large tracts of old growth forest, as I think it was, the reintroduction of the beaver may have been central to its persistence. If this hypothesis is valid, there is considerably more potential habitat today than there was in Tanner’s era; much of this potential habitat has been overlooked or dismissed in organized search efforts; and the dismissals of post-Tanner reports based on his habitat model rely, at least in part, on a false premise.

 

 

 


Scaling Data 2012-2016

To expand on some of the data included toward the end of the March trip report (which is worth reading in in conjunction with this post), I thought it would be informative to provide a season by season and sector by sector breakdown of the scaling I and others involved with Project Coyote have found since the spring of 2012. To do so, I’ve gone through my notes and photographs and have done my best to reconstruct the data collected. While not complete (I’m quite sure a good deal more scaling was found in Sector 3 during 2013-2014, for example), I think this breakdown is a fairly accurate reflection of what we’ve found over the years.

As discussed in previous posts, I think extensive scaling on hickory boles is the most compelling for Ivory-billed Woodpecker. Bark on this species is thick, dense, and usually remains very tight for a long time. Extensive scaling on sweet gum boles and oaks (upper boles and large branches) is second among work that I’ve found. Work on small boles, and higher and smaller branches is somewhat less compelling and is more significant for its abundance. Some of the high branch scaling and work on smaller boled sweet gums may well have been done by Pileated Woodpeckers (and possibly by Hairy Woodpeckers), but the abundance, the presence of large bark chips in many cases, the way it appears in clusters, and the fact that Pileateds scale infrequently suggest a different source for much of it.

I have excluded all work where squirrels are suspected but have counted one tree, a hickory found this year, on which the work could well have been that of a Hairy Woodpecker. Hairies do forage for Cerambycid beetles just under the bark, but they’re only capable of removing tight bark in small pieces; their work on hickories is perhaps more accurately described as excavation through the bark.

The trail cam images toward the end of this post are the best we have (out of many thousands of hours of coverage) showing how these species forage on suspected ivorybill feeding trees.

All trees were live or recently dead (twigs and sometimes leaves attached). All scaling was on live or recently dead wood.
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Totals

Sweet Gum (Liquidambar styracifula)

Sector 1:         46

Sector 2:         8

Sector 3:         51

                        105         (84.68%)        

~15% had scaling on boles (a few of these were large trees). The majority of work was on crowns, including larger branches. Fallen trees were included when woodpecker involvement was evident and bark was tight.

Bitternut Hickory (Carya cordiformis)

Sector 1:            3

Sector 2:            4

Sector 3:            7

                           14         (11.29%)

All trees were standing; scaling was on boles and was very extensive (the tree shown on the homepage is one example) with one exception from this year . Insect tunnels were visible in all examples. An additional hickory with a modest amount of high branch scaling was found in Sector 1 this year but was not counted for this analysis.

Oak (Quercus) spp.

Sector 1:         1

Sector 2:         4

Sector 3:         0

                         5         (4.03%)

All oaks had scaling on large branches; one also had some on the bole. All oaks in Sector 2 were found in a single cluster.

We have some information on forest composition in Sector 3, and it appears that sweet gums make up approximately 19%, oaks upwards of 35%, and hickories somewhere under 10%. Sectors 1 and 2 may differ and be more varied in overall composition.

The overwhelming preference for sweet gums relative to their abundance stands out. The scaled oaks are a mix of species, one Nuttall’s, one willow, the others unidentified.

In Sector 3, I am treating the compact stretch from the location of Frank Wiley’s sighting last spring/downed sweet gum top where we had the camera trap to just south of our current deployment as a cluster. The estimate of 23 trees being found in this area is conservative. I have only found one instance of recent scaling north of the location of the downed limb/Frank’s 2015 sighting. The main cluster has been in the same vicinity this year and last, with additional work scattered around farther south. Two of the hickories are within 30 yards of each other, approximately half a mile from the cluster, and one was on the edge of the concentration.

It also may be significant to note that we found a cluster of old but intriguing cavities in the same vicinity as the Sector 3 concentration in 2013-2014. Most of these seem to have fallen. The difficulty we’re having finding active, suggestive cavities is vexing, and may be the most compelling reason to be skeptical about the presence of ivorybills in the area. At the same time, finding Pileated cavities is difficult, even in defended home ranges.

I’m treating Sector 1 as a single concentration; the vast majority of the work is on a natural levee where sweet gums are abundant. The entire area is considerably larger than the other clusters, but given the abundance and ease with which we’ve found sign there over the last five seasons, I think it constitutes one area of concentration.

In Sector 2, there was a small cluster in the area where I recorded putative kent calls in 2013, with work found in 2012 (spring and fall) and 2013. Because the area is small with open sight lines, I can be confident there has been no recent work there since late in 2013 (I last passed through it with Tom Foti back in March of this year.)

The sweet gum work Tom and I found on that day was perhaps half a mile north of this cluster, within 100 yards of the hickory on the homepage. The other hickories found in the 2013 and 2014 seasons were not far away, no more than 500 yards apart as the crow flies.

There’s obviously some bias here, since there’s a relationship between finding feeding sign in a given area and spending time there. Nevertheless, I have little doubt that the putative ivorybill work tends to be clustered. I also have little doubt about the strong preference for sweet gums, since I’m not looking at tree species when I look for scaling. The degree to which sweet gums are favored has only become clear over the last year or so.

Frank pointed out this data does not reflect most of the scaling that likely exists in relatively close proximity to the Sector 3 cluster but cannot be quantified because it is in an area we have intermittently visited due to  inaccessibility. Only two or three examples are from this area, which has been visited a handful of times.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 


Old Material, New Light: More from the Archives Part 2

In late December 2014, I wrote what I’ve described as a speculative post titled, “Is There a Way to Recognize Ivory-billed Woodpecker Excavation? In that post, I relied on Tanner’s Plate 11,

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Tanner’s Plate 11, “Dead hackberry, fed upon frequently by Ivory-bills”. Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

a brief description from the monograph: “When Ivory-bills dig, they chisel into the sap and heartwood for borers like other woodpeckers, digging slightly conical holes that are usually circular in cross section (Plate 11)”, and online imagery showing the work of other Campephilus woodpeckers. Material found during my recent visits to Kroch library at Cornell lends some support to the ideas contained in that post, and so does T. Gilbert Pearson’s photograph of a tree that had been fed on by ivorybills. Holt:Pearson Tree

The archival material includes additional images of ivorybill excavation and a considerably more detailed description by Tanner in a document prepared for the Cuban search in the 1980s. The passage includes somewhat more detail on bark scaling than is found elsewhere, but more importantly it describes ivorybill excavations as “hard to distinguish from similar digging by the Red-bellied Woodpecker”.

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Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

This description may seem counterintuitive to some. Despite my own writing to the effect that ivorybill morphology may lead the species to dig less efficiently than pileateds and my references to targeted digging, I still had an underlying assumption that the size of the bird would correlate with the size of the dig and that ivorybill excavation would often resemble the familiar large furrows dug by PIWOs. While a couple of the holes in Plate 11 and in Pearson’s photograph may well involve the merging of more than one dig, it appears that ivorybill excavations are usually more targeted and that large furrows are not typical.

Also of interest for multiple reasons, including the observation of birds scaling very small limbs and of one feeding 5′ from the ground, are Tanner’s field notes from April 3rd, 1937.

IMG_1100

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Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

I’ll let the remaining images of known and suspected ivorybill excavations speak for themselves and will conclude with a few from our search area that seem consistent with known ivorybill work. While I’m nowhere near as confident about this material as I am about scaling, I suspect that finding excavations that are consistent with what ivorybills are known to have done in conjunction with scaling is suggestive.

I hope this material will be useful for other searchers. All images from the Singer Tract below are courtesy of the Division of Rare and Manuscript Collections, Cornell University Library. Most of these images were published in Tanner’s dissertation but have not been widely disseminated.

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OakNest copy

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1935 Nest Tree, Red Maple

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1935 Nest Tree and Detail from a Different Perspective

And now some examples from our search area that resemble the existing images of known ivorybill excavation. This is not something I’ve focused on, so I’ve probably missed other examples.

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Image at bottom is a detail of the sweet gum snag above. I suspect that more than one species of woodpecker is involved.

There will be one or two more installments in this series, but the next post is likely to be a trip report, probably the last for this season.


Strips, Flakes, Chips, Chunks, and Slabs: Squirrels, Pileated Woodpeckers, and Ivorybills, Part 4

Careful examination of bark chips found in conjunction with extensive scaling is one of the key elements in our diagnostic gestalt, but “chips”, a term I’ve been using for years, is both inaccurate and too vague for what we believe is being left behind by Ivory-billed Woodpeckers and for differentiating it from the leavings of other animals. Tanner used “pieces” of bark, ranging “from the size of a “silver dollar to the size of “a man’s hand.” A caption from the National Geographic article on the 1935 Allen and Kellogg expedition that refers to “large chunks of bark”.  The existing images of these pieces of bark suggest that chunks is the better term.

It’s important to reiterate that this discussion applies only to live and freshly dead hardwoods. Pines slough bark quickly after death. The process is slower in hardwoods, but as decay progresses, the bark loosens considerably, with the rate of loosening depending on species and environmental conditions. Once the bark has loosened sufficiently, PIWOs can and do scale bark extensively, sometimes leaving behind large chips. In the images that follow (from Allen and Kellogg and Tanner), the bark chips ascribed to ivorybills appear to come from considerably longer dead trees than some of the examples we’ve found, but the images are informative.

Ivorybill Scaling Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

Ivorybill Scaling Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

The small tree shown above, identified as a “dead gum” by the 1935 expedition, appears to be a hackberry or sugarberry not a gum, and a fairly long dead one; the pieces of bark at the base resemble ones we found beneath hackberries or sugarberries in our old search area, some of which were considerably larger (the one below is the largest).

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This colorized slide reveals more about the bark at the base of these pines than the black and white print in Tanner (Plate 9).

There’s also this example, (Plate 10 in Tanner), which appears to be in a considerably more advanced state of decay, and presumably looser, than much of the work we find most interesting. I suspect most of the grubs were placed on the chip for illustrative purposes; the caption “Beetle larvae from beneath bark of Nuttall’s oak” is ambiguous as to where the larvae, which appear to be small Cerambycidae, were actually found.

What I think is most salient in Tanner’s description of bark chips is shape not size. In this regard, it seems important to come up with a more specific set of terms to replace the commonly used “chips”. I’d suggest using chunks and slabs for suspected ivorybill work (although smaller pieces of bark may also be present). Pileated bark removal can involve chips, strips, or flakes, the last when they’re doing the layered scaling discussed here and here. I suspect that squirrels remove hardwood bark primarily or exclusively in strips, and of course, their bark removal on cypresses leaves shredded bark hanging from the trees.

Let’s take a closer look at the differences among pieces of bark we have reason to believe were left by squirrels, those we have reason to believe were left behind by Pileated Woodpeckers, and those we suspect were left behind by Ivory-billed Woodpeckers.

I collected a number of bark chips from the tree we know to have been scaled by a squirrel, and while these were removed before our camera trap revealed the source, there’s strong reason to think they too were left behind by squirrels.

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Note the uniformly elongated shape and the ragged appearance at the tops and bottoms of these strips of bark. This is not typical of bark that we infer or know to have been removed by woodpeckers, and it’s consistent with chewing, not scaling. The presumed squirrel strips I collected had the following dimensions:

9”x2.5”

7”x2.25”

5.75”x2″

7.5”x1.75”

4”x1.75”

The downed sweet gum from which they had been removed was a fairly young tree, and the bark is much thinner than on more mature ones. These strips were approximately 1/8″ thick. While this is a very small sample, we suspect (along with Houston from IBWO.net) that approximately 3″ is the upper limit for width when a squirrel is doing the bark removal.

Our research and observations suggest that Pileated Woodpeckers have two strategies for removing tight bark; one involves pecking around the edges until they can gradually pry off small pieces, and the other involves scaling away strips, sometimes in layers. Their physical structure precludes them from doing the extensive, clean scaling of tight bark that Tanner associated with ivorybills.

We suspect that this collection of chips, from a honey locust near a known Pileated nest, reflects the range of what the species is capable of doing on a tight-barked hardwood (and honey locust bark is relatively thin). The upper limit appears to be hand size, with many-quarter sized or smaller.

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The following are measurements of some fairly typical suspected Pileated strips from a sweet gum:

7”x1”

8”x.8”

7”x.8”

6”x.8”

The strips shown below, suspected Pileated Woodpecker leavings from a high branch, are on the large end of the spectrum for this category of work. The Peterson Guide is 9.5″ x 6.5″. I can’t rule squirrel out completely for these.

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Flakes resemble strips, but they are removed in layers, so that reaching the sapwood is a gradual process. Pileated scaling frequently has this appearance, something that seems frequently to be the case with congeners, including the larger-billed Black Woodpecker (Dryocopus martius).

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The chunks and slabs we suspect to be ivorybill work are significantly larger and thicker than strips, flakes and chips, although strips and chips may be present in the mix at the base of suspected feeding trees. Chunks are usually more irregular and varied in size and shape, and both chunks and slabs sometimes have what appear to be strike marks from a broad bill.

I kept one of the chunks scaled from the hickory tree on the homepage, a fairly typical example. It is 8.5″x3.5″ and .375″ thick. (It has undoubtedly lost some of its thickness after drying for over two years.)

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The sweet gum chunk with the apparent bill mark Frank is holding is 7.5″x3″ and .25″ thick. On mature, thicker barked trees most or all suspected ivorybill chunks, chips, and slabs will have been removed cleanly, all the way down to the sapwood.

Frank adds:

This particular bark “chunk” is intriguing on several levels. We have found that markings many describe as “bill marks” are really truncated galleries between the bark and sapwood. Marks made by woodpecker bills are distinctive, but somewhat subtle, and easily overlooked. This chunk actually has two interesting markings – markings that were left by the animal that removed the bark. The first is near the end of my left thumb – my right index finger is pointing toward it. It is about a quarter inch wide, a bit over a half inch long, and three sixteenths of an inch or so thick. The other is a “V” shaped “notch” at the end of the chunk, near the center of the photo. These places look as if they’ve been struck with a chisel – hard enough to rip the bark away from the sapwood/cambium. This suggests that, even though this bark was very tight, very few strikes were required to loosen and remove it. Granted that these marks are bill strikes, this suggests that the bird removing bark is indeed a powerful animal for its size. Back to Mark.

DSC00031The two preceding examples are on the smaller side for suspected ivorybill work; in the first, the density, tightness, and grain of hickory bark seem to be a limiting factor on size. Some of the larger examples are shown in the Bark Chip Gallery (as are several of the images shown above). A couple of additional examples of larger slabs are below. In the first, the oak was approximately 8 months dead (leaves attached), and the bark was still tight. (The fractured slab was damaged in transit.)

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Scaling and Squirrels, Part 3, A Closer Look at Surfaces and Edges

The first two installments in this series, which was inspired by the discovery that squirrels are doing some of the bark scaling in our search area, are here and here. This installment will consider the appearance of the scaling itself, and the next will focus on pieces of bark; “chips” no longer feels sufficient to describe the spectrum of what we’ve found, and using more specific terminology may shed more light on what we think is diagnostic and why.

I’ll begin with the work we are now presuming to have been done by squirrels. In retrospect, it’s easy for me to understand why I was fooled by this bark stripping. I was not seriously considering mammals as a source due in part to the extent of some of the work involved; I never saw incisor marks on the wood, something that’s often described as being an important indicator; similarly, I have been unable to find these marks in the photographs I’ve taken of stripping that we now presume to have been done by squirrels.

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Tanner’s Plate 8, “Ivory-bill feeding sign on slender limb. Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

In addition, I was somewhat misled by Plate 8 and the description of what Kuhn thought was diagnostic that his daughter shared with us. (Scroll down in this post to see where my thinking went astray.) It’s my current view that Plate 8 could conceivably be squirrel work. Tanner doesn’t state that he actually observed an ivorybill doing the scaling.

Between 1937 and 1939, Tanner observed actual scaling a total of 73 times, but it’s not clear how many instances he might have photographed. In hindsight, the scaling in Plate 8 is not particularly impressive; the scaled patch is relatively small; the bark is thin and the hanging pieces may be an indication of removal by gnawing rather than bill strikes. Adhering bits of shredded bark and cambium are evident in some of the work we believe to be squirrel, including on the tree where we captured a squirrel stripping bark (albeit much smaller ones in that case).

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Small bits of bark and cambium adhering to downed sweet gum limb after presumed scaling by squirrels.

In fact, I think one of the keys to recognizing that squirrels are likely responsible for removing bark is a ragged, shredded, or chewed up appearance to the bark and cambium, as in the examples below.

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The information we got from Mrs. Edith notwithstanding, I am going to examine the edges of scaled areas more carefully and be sure they are for the most part cleanly incised. This is one of the main criteria for ascribing the work to ivorybills (although by no means all Campephilus scaling falls into this category, and some can have ragged edges). As I’ll be discussing in the next post, I think that bark chip characteristics provide an even better diagnostic.

Now let’s turn to targeted digging and the similarities between what we’re finding and the work of other Campephilus woodpeckers. In some of my previous posts on bark scaling I’ve mentioned “little or no damage to the underlying wood”. “Little” is the operative word here. In most if not all of the examples of scaling associated with large Cerambycid exit tunnels that we’ve been able to examine up close, there are also indications of targeted digging, and we have seen similar targeted digging on some of the higher branch work we’ve found. Targeted digging involves the expansion of individual exit tunnels in varying degrees. This can range from what appears to be little more than probing with the bill to deeper and wider excavations, but this excavation is incidental to the scaling, whereas in Pileated Woodpeckers, scaling on tight barked-trees is typically incidental to excavation.

A magnificent series of photographs by Luiz Vassoler posted to the Flickr Campephilus group, showing a Crimson-crested Woodpecker scaling and doing targeted digging, is illustrative (scroll through to your right for the whole series). This is not to suggest that other woodpeckers can’t or don’t dig for larvae in a targeted way, only it’s more suggestive evidence for the presence of Ivory-billed Woodpecker in our search area, given the context and what’s known about the foraging behavior of  its congeners.

I’ll keep commentary to a minimum and post some examples from our area (the seven photos immediately below) and then links to work done by other Campephilus woodpeckers.

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Pale-billed (on palm):There’s no scaling here, but the exit tunnels have been expanded vertically, and the expansions resemble some of the rectangular ones above.

Pale-billed (at nest): Very targeted digging and slight expansion of some exit tunnels.

Pale-billed: On a scaled surface, some tunnels expanded.

Robust: I’m including this almost as much for how well it shows Campephilus foot structure and the rotation of the fourth toe and hallux.

Cream-backed: Somewhat more aggressive expansion of tunnels on longer dead wood.

Red-necked: Targeted digging to the right of the bird.

Crimson-crested: Elongated dig into exit tunnel.

Crimson-crested: another great shot of the Campephilus foot. Targeted dig at bottom of scaled area.

Crimson-crested: Video showing targeted digging on an unscaled area.

Magellanic: are the most Dryocopus-like in the genus in terms of foraging behavior. Note their smaller bills and relatively shorter necks. They seem to spend a lot more time feeding near the ground and excavating large foraging pits than the other species, but they too do a considerable amount of targeted digging.

Magellanic: The appearance of the scaling here is strikingly similar to what we think is diagnostic for ivorybill. The tunnel at bottom right has been expanded, likely with the tip of the bill. It looks as though there is more targeted digging above and to the left.

The two images below, showing targeted excavations on small limbs, associated with extensive scaling on young, freshly ambrosia beetle-killed sweet gums, bear a striking similarity to work by Crimson-crested  (the long furrows in particular) and Magellanic Woodpeckers.

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One final and more speculative observation that might be of interest to other searchers. I had dismissed this work on a live maple as likely pileated because it is generalized digging, not scaling. After going through so many images of Campephilus foraging sign, I’m a little more intrigued by it, as I see similarities to sign like this and this. Like some of the work on ambrosia beetle-killed sweet gums, this almost looks like a hatchet had been taken to the tree; the wood was not at all punky; and red maple at 950 on the Janka hardness scale, while not nearly as hard as bitternut hickory (1500), is harder than sweet gum (850). While I’m not proposing this as a diagnostic, it may be more interesting than I initially thought.

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Unusual excavation on live maple

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Scaling and Squirrels: Part 2, Digging Deeper

Part 1 of this series is here, and the event that led to my writing it is discussed here.  I now expect to write 2-3 additional posts on this topic and may create a new page that summarizes the whole series. I’ve hidden the Bark Scaling Gallery page to be reworked later or incorporated into the summary.

This post will reiterate, revise, and expand upon earlier ones dealing with bark scaling and woodpecker anatomy. The next one will focus on certain characteristics of the scaling we think is being done by Ivory-billed Woodpeckers, on finer details that characterize it (based in part on comparison with work done by congeners), and on how to differentiate it from bark removal done by squirrels. The following entry will deal with bark chips in more depth, and from a slightly different angle than previous posts on that subject.

I had originally intended to address the next post’s planned content in this one, but as I started writing, I realized the long but necessary introduction would bury the lede. It soon became clear that I’d have to divide the post in two with this one for background.

The first important point is that woodpecker taxonomy is in a state of dramatic change, so much so that the American Ornithological Union is being advised to place Downy and Hairy Woodpeckers in separate genera and that their current genus, Picoides, should be divided into four. Notwithstanding the taxonomic upheaval, there’s no question that Campephilus woodpeckers and Dryocopus woodpeckers are only distantly related, that their similarities are the product of convergent evolution, and that these similarities are far more superficial – involving size and coloration – than structural or behavioral. Formerly, some incorrect taxonomic assumptions led to the lumping of Campephilus and Dryocopus into the “tribe” Camphelini, an idea that’s discussed and dismissed in the first paper linked to above. This has been one factor in perpetuating some fairly common and persistent misconceptions – that the two species are closely related, that they occupy or occupied the same ecological niche and might be competitors, and that hybridization might be possible (something I hear surprisingly often).

The following differences are relevant to this discussion:

  1. Bill size and shape. These are dramatically divergent as any comparison shot of specimens makes clear. It’s also worth noting that the three North American Campephili are closely related to each other. DNA analysis suggests the three are distinct species and the Cuban ivorybill may be more closely related to the Imperial Woodpecker than the mainland US species. This study suggested that divergence among the three took place between .08 and 1.6 million years ago. The southern members of the genus are more remote cousins, having diverged approximately 3.9 million years ago. At one time, the southern species were considered a distinct genus, and they have smaller bills, both objectively and relative to body size. Magellanic Woodpeckers have the smallest bills relative to body size in the genus, and their foraging behavior is more Dryocopus-like than their congeners’. DSC00866
  2. Neck length. The much longer neck of the ivorybill allows for a broader range of motion.
  3. Foot and leg structure. Campephilus woodpeckers have a unique variation on what have been called pamprodactylous feet. (Wikipedia and David Sibley both miss the vast difference between Campephilus foot structure and that of most other woodpeckers.) In this genus, the hallux (first) and fourth toe (the rear toes) are both on the outer edge of the foot; the toes can be rolled forward for climbing and backward for perching in a manner that looks more zygodactylous. (The preceding links to images of Sonny Boy, the juvenile ivorybill, and Kuhn are great illustrations.) The fourth toe is highly elongated, the longest toe on the foot, and the hallux, (in the ivorybill, the outermost toe) is relatively longer than in any climbing woodpecker species. The second and third (innermost toes) are angled inward. This is shown quite clearly in a number of the images from the Singer Tract, including Plate 13 in Tanner.

    Enlargement of image used for Tanner's Plate 13 showing foot structure. Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

    Detail of Tanner’s Plate 13 showing foot structure. Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

  4. Dryocopus woodpecker feet are closer to being truly zygodactylous – two in front, two behind, with limited mobility and the hallux as the inner rear toe, although the fourth toe can be rolled outward to some extent; this provides less stability when making lateral blows.
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    Pileated Woodpecker foot showing zygodactylous structure and slight outward rotation of fourth toe. Photo courtesy of Carrie Griffis, who posted it on the Woodpeckers of the World Facebook group and kindly granted permission to include it here.

    In addition, Campephilus woodpeckers typically climb and forage with their legs both farther apart and higher relative to their bodies than Dryocopus. This enables them to keep their lower bodies closer to the trunk and move their upper bodies more freely, providing more stability for making powerful, lateral blows.

    4. Tail structure: the ivorybill’s tail feathers are long, thin, barb-like, and stiffer than the pileated’s. The tail serves as an anchor and also helps allow for a broader range of motion.

    Tailfeathers

    Middle Tail Feathers: Flicker, Ivory-billed, and Hairy Woodpecker

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    Pileated Woodpecker Tail Feathers. Note how the longest one resembles that of a Flicker more than that of an ivorybill.

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    5. There other structural differences, including wing shape, but these are the main ones that point to how Ivory-billed Woodpeckers have evolved in a way that makes bark scaling their most efficient foraging modality, whereas Pileateds are far better suited to digging, using a perpendicular motion.

Much of the foregoing is based on Walter Bock’s  analysis of woodpecker adaptations for climbing, which was also discussed in depth here. I’ve tried to explain Bock’s key points in straightforward and less technical terms. A longer quote from Bock appears at the end of this post.*

In addition to these structural differences, Pileated Woodpeckers (and to the best of my knowledge all their congeners) regurgitate when feeding young. Campephilus woodpeckers carry food to the nest and appear to be highly dependent on beetle larvae when caring for their nestlings. This means that Pileated Woodpeckers have to ability to take advantage of multiple food sources during nesting season, while Ivory-bills have a more limited range of options. While I don’t think this supports Tanner’s theory of old-growth dependence, it does point to a higher degree of specialization that would impact numbers, range, and suitability of habitat.

At the same time, the anatomical differences and degree of specialization convince me that certain types of feeding sign are beyond the physical capacity of a Pileated Woodpecker and are likely diagnostic for Ivory-billed Woodpecker.

There is a dearth of clear images showing Ivory-billed Woodpecker feeding sign. There are a handful of photographs, most of them very poor. The majority were taken in the Singer Tract and some showing work on pines were taken in Florida by Allen and Kellogg.  Few of them depict the high branch work that Tanner described as being characteristic, and when they do, there’s virtually nothing that can be discerned from them. It is also not entirely clear that Tanner’s attribution of feeding sign to ivorybills was always based on direct observation, which makes us wonder whether some of the work might actually have been done by squirrels. Regardless, this makes it difficult to draw inferences from the existing body of imagery.

That said and with awareness of the perils in extrapolating, one lesser known image from the Singer Tract is worth comparing with the work on boles that’s been discussed in multiple posts.

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“The Blind at Elm Rock”, Ivory-billed Woodpecker nest tree and detail showing scaling and excavation on trunk. Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

This is a view of the 1935 nest tree, which was a red maple. It’s taken at a different angle than the more familiar shots, so it shows some large areas of scaling on the bole that the others do not. While I can do no more than infer that this was done by ivorybills, it’s clearly old, and there’s an abundance of excavation in the underlying wood; nevertheless, the edges and contours of the scaling are strikingly similar to the work we’ve found on boles, especially the area at the lower right, just above the intervening foliage.

This is the jagged appearance I described in the previous post; the similarities are most evident in the picture below and on the home page. ScalingNewArea

Because there are so few informative images of ivorybill feeding sign, the best available option is to look at the work of other Campephilus woodpeckers. Even though they are not as closely related as the Cuban ivorybill or the imperial, their morphology and foraging behaviors are similar; even the work of the smaller-billed but oft-photographed magellanic can provide some clues. I’ll examine this and some probable identifying features of squirrel scaling in the next post, which will take a close look at scaled patches on trees.

*”. . . in most woodpeckers, as, for example, the pileated woodpecker, the legs are held more or less beneath the body,the joints are doubled up,and the tarsus is held away from the tree trunk. This position of the legs is disadvantageous for the bird, because the body is held away from the tree trunk and the muscles of the leg are working at a mechanical disadvantage; the analogy is to the mountain climber who is standing on a narrow ledge with hand holds only beneath his chest. In the ivory-billed woodpecker, the legs are directed away from the center of the body, and the tarsus is pressed against the tree trunk. This method allows the body to be held close to the tree, with the joints of the leg extended. Hence the leg muscles have a mechanical advantage, because they are at the beginning of their contraction cycle and are acting along the length of the segments of the leg. When the body is held close to the trunk, it not only decreases the outward component of gravity but allows the tail feathers to be applied to the supporting surface for a greater distance from their tips. If the bird is climbing on smaller limbs, the feet can encircle the limb and thus obtain better support. However, no matter what size the limb is, the disposition of the legs and the spreading of the toes of the ivory-billed woodpecker furnish direct and powerful resistance to both the lateral and backward motions of the woodpecker when it is at work and, with the tail, furnish a tripodal base of great strength against the pull of gravity.”