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.


Bits and Pieces Part 3: Internalized Beliefs, How They Got That Way, and What the Record Really Shows

Part 1 is here. Part 2 is here. It looks like this series will end up being a five-parter. Part 4 should follow later this week and Part 5 sometime thereafter.

Most of us with an interest in the ivorybill have internalized a set of beliefs about what constitutes “suitable” habitat. To a large extent these beliefs treat the Singer Tract as a model – a vast tract of “virgin” bottomland forest dominated by oaks and sweetgums, with abundant, moss-draped cypress for atmosphere (although ivorybills seem to have avoided cypress in the Tract). The habitat description in Stephen A. Shunk’s excellent Peterson Reference Guide to Woodpeckers of North America clearly expresses some of these beliefs, which have influenced the overwhelming majority of modern search efforts and which are embedded in the minds of most searchers (myself included) to the point of being a default:

Virgin bottomland forest almost always below 100 ft. (30 m) elevation. May also have occurred in uplands but by 1900 restricted to areas downstream of pine-bald cypress interface. Requires large tracts of contiguous forest with very large-diameter trees and adequate dead and dying trees to provide forage and nest sites.

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The Guide, which draws on an 1891 article by Hasbrouck for the 100 foot elevation, goes on to list three different habitat types described by Tanner – sweet gum-oak dominated forests in the southeast (with species of oak varying depending on location) outside of Florida; river swamps in Florida dominated by cypress, black gum, and green ash; and creek swamps in Florida characterized by cypress, red maple, laurel oak, black gum and cabbage palmetto, with feeding in adjacent pine woods.

This is not to criticize the Guide or its author – I recommend the book highly and the overall treatment of the ivorybill is thorough and evenhanded. The quote is intended to point out the pervasiveness of these ideas about habitat requirements, ideas that Tanner reinforced, especially in later years. They’re so pervasive in part because the myth of the “virgin forest” has shaped ivorybill lore since well before Hasbrouck and has influenced almost all habitat assessments since Tanner.

The virgin forest myth is a topic for another day; the central point is that ivorybills have been found in more diverse habitat types than most have believed. It’s worth bearing in mind that Tanner himself asserted that ” . . . at present the only suitable habitat for ivorybills is in tracts or areas of virgin timber”, a narrow, almost lawyerly, and largely conjectural conclusion – one not entirely supported by fact. Ivorybills bred in at least one Singer Tract area (Mack’s Bayou) that was predominantly regrowth, and as has been discussed in previous posts, Tanner became more dogmatic and blinkered about habitat requirements in later years, dismissing the John Dennis Texas recording because a Pine Warbler was captured on the tape.

As should become clear, Ivory-billed Woodpeckers were found in a variety of different habitats and did not always require extensive tracts of contiguous forest. Large diameter trees, their importance, and what Tanner meant by “large” are discussed in another post, but even if one accepts that large trees were preferred in the Singer Tract, much of the feeding sign Allen and Kellogg documented in Florida was on small, fire-killed pines.

My plan for this post was to focus on Ivory-billed Woodpecker records from outside the historic range as delineated by Tanner in his oft-reprinted 1942 map and adaptations thereof.

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US Fish and Wildlife Service range map based on Tanner, including locations of selected post- Singer Tract reports

The purpose behind this original plan was in part to show that the historic range of the ivorybill was considerably more extensive than is commonly believed, but more importantly to show that ivorybills inhabited more varied habitats than is commonly believed. As time went on, my focus has shifted even more toward the question of habitat diversity, though the fact remains that the historic range was considerably more extensive than the Tanner map, or the one drawn by Hasbrouck in the 1890s, might lead one to expect.

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Hasbrouck’s Ivory-billed Woodpecker Range Map (1890)

In recent Facebook comments, several ornithologists have suggested that without physical evidence, no record should be accepted. As I see it, this standard is, to some extent, a kind of ahistorical overkill, since it has never been applied in the past. Post-1939 Singer Tract observations don’t meet it, so the last record date is pushed back by several years, which can impact statistical analyses. At the same time, the parsimonious approach is not entirely meritless, since it eliminates false positives. The problem is that there’s no purely objective standard for evaluation of historic (and pre-contact) records, even when it comes to specimens. In many cases, location information for specimens is non-existent or ambiguous; for example, one specimen in Cornell’s collection (1896) is listed as coming from the “Florida Keys”; “Key” in this context more likely refers to an island of forest surrounded by the Everglades than to the islands offshore.

Thus, in this post and the next, I’ll be looking at many reports from within the recognized historic range, as well as some from beyond those boundaries. I will be focusing on reports accompanied by physical evidence or published accounts stating that physical evidence was obtained but will include or mention a few additional ones that seem particularly credible based on the source or amount of detail.

The more surprising of these reports are unsupported by physical evidence. These come from the Eastern Shore of Maryland, Swedesboro, New Jersey, and Pennsylvania. Most date to the 18th-century, a time when ivorybills were reported to have fed on trees girdled for clearing. The most interesting of these come from Peter Kalm, a Swedish naturalist and student of Linnaeus. They have been discussed in several articles by Benjamin Leese, who has also written about early records from Ohio, Indiana, and Kentucky (for which the case is stronger). Most of these papers do not seem to be available online.

Just under 200 of the 418 specimens (including Cuban ivorybills) listed by Hahn provide no location information or merely identify the state, or country in the case of Cuba, where the collection took place. There are multiple cases in which specimens were reportedly collected but have not been found, and in several cases, there’s no way to correlate the claim of collection with an actual specimen. In one instance, not catalogued by Hahn, a pair of ivorybill specimens was mislabeled as Pileated Woodpecker until the error was corrected in the 1960s.

When it comes to pre-Columbian sites, there’s no way to be certain whether material collected from graves and middens involves trade goods or locally killed animals, although tarsometatarsi are likely local, especially east of the Mississippi, where there’s little evidence to suggest that woodpeckers had ceremonial value. Leese addresses this subject in a paper on Native American uses of ivorybill parts. I have included records involving tarsometatarsi and one from a West Virginia midden that involves parts of two lower mandibles.

I’ve created a google map showing the locations of the records from habitat types that don’t fit the ‘large tracts of contiguous bottomland forest’ paradigm. I’ve provided some details for each location. The map draws on Appendix E or the US Fish and Wildlife Service’s Recovery Plan, Tanner, and Jackson. I’ll discuss the records from coastal areas, south Florida, the upper Mississippi and locations east of the river in the next post. I’ll conclude this one with a look at the records from the western edge of the range, since they relate to the Nebraska report and why I thought it might be credible; the Nebraska location is actually somewhat east of the records from the southern plains.

One record that I find compelling does not include a specimen; it’s from the 1820 Long Expedition, the first scientific exploration of the American West. Screen Shot 2017-10-08 at 8.08.14 AM

The ivorybill’s call is described, and Pileated Woodpecker is distinguished and described as common in the area. These facts lend credibility to the report, as does the fact that Thomas Say was the expedition’s naturalist. While Say is best known as an entomologist, the expedition produced the first descriptions of a number of bird species, and an entire genus of flycatchers was named in his honor.

The location of this record is approximately south of Tulsa on the Canadian River, near the 96th Meridian. This is farther west than the location of the erroneous southeastern Nebraska report and well into the eastern Great Plains. It is also well beyond the range of the bald cypress. The relatively narrow floodplain would have been dominated by cottonwoods and willows, as it is to this day. This image, from ca. 1920s shows “Standing Rock” a geological feature now flooded that was discovered by the expedition a day after the ivorybill encounter. It should add a visceral sense of the area’s appearance to supplement the description above.

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Even if one opts to reject this record for lack of physical evidence, there are several others from approximately the same longitude that do involve specimens, and some are from the 20th century.

The Recovery Plan suggests that there are two records from west of Tulsa, a specimen was “probably” collected by Woodhouse along the Cimarron River, Pawnee County in 1849. Per Jackson, the specimen was sent to the Philadelphia Academy of Natural Sciences, which has four specimens without location information in its collection. The second is from House Creek in Pawnee County, also Woodhouse 1849, and also reportedly sent to the Philadelphia Academy but not found there.

There are several Plains records from Texas. A specimen currently in the Dallas Museum was collected on Bois d’Arc Island, just southeast of Dallas, elevation 400′, in 1900. There were multiple reports from the area through 1910, and an additional bird may have been collected in 1918. A bird was reportedly “caught in a trap” in nearby Kaufman County in 1927 and examined by an R.E. Huck but not preserved. An additional Texas record, from farther south but west of the 96th Meridian, comes from New Braunfels County, south of Austin and east of San Antonio. There were multiple reports ca. 1900, with a collection reported but no specimen preserved.

Although only one record from the eastern Plains can be attached with certainty to a currently existing museum specimen, there’s proof that Ivory-billed Woodpeckers were found in this region as recently as 1900 and considerable circumstantial evidence for their presence along riparian corridors on the plains of Texas, and possibly Oklahoma,  into the 20th-century. The habitat involved is markedly different from what so many have believed ivorybills require. I’m not suggesting that ivorybills persist at the western edges of their historic range, but as will be discussed in the final installment of this series, I think their ability to exploit these relatively narrow, willow and cottonwood dominated floodplains can help explain how the species could have persisted into the 21st century.

Stay tuned for Part 4.


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.

 

 

 


Not So ‘Virgin’ Forest: The Singer Tract Myth Debunked

Update: This post includes hard data about the extent of old growth in the Singer Tract (scroll down past all photographs) and in ivorybill home ranges. The general points made below remain valid, although some of the wording is perhaps too strong; Tanner overestimated the amount of old growth in the Tract ( at “over 80%” v. 72% in fact), and the Mack’s Bayou home range was predominantly second growth.

This post is a companion to the previous one and to others discussing habitat conditions in the Singer Tract. Those posts reference Richard Pough’s 1944 report to the Audubon Society. Pough, whose study was never published, noted that much of the Singer Tract had actually been under cultivation prior to the Civil War. But it’s worth taking a closer look at just how much.

Tanner characterized the Singer Tract as “the largest tract of virgin timber in the Mississippi Delta,” contending that it contained “120 square miles of virgin forest in 1934”. He also wrote that the largest plantation “had about 3000 acres under cultivation,” while suggesting that “some of the early settlers along the Tensas River cleared land along the river banks for cotton fields.” Thus, the myth of the Singer Tract as virgin forest was born.

It’s not clear where Tanner got his information, but some of his characterizations are not supported by the historical record; the language about “early settlers” almost seems disingenuous when one looks at the history of Madison Parish. (Rootsweb has many pages devoted to this subject, and I’ve drawn heavily on them for this analysis.) As should become evident, there was a great deal of human activity in and around the Singer Tract, especially prior to the Civil War. I will suggest that most, perhaps virtually all, of the arable land in the parish, had been cleared for agricultural purposes and that the Singer Tract was a mix of second growth and remnant old growth, most of which was in the lower-lying, wetter areas that Tanner deemed to be less suitable for ivorybills.

A 1937 Masters thesis in economics by Robert L. Moncrief, “The Economic Development of the Tallulah Territory”, provides a great deal of information about the parish and its history. In the post-Columbian era, the area was very sparsely settled until the 1830s. Madison Parish was established in 1839, and in 1840, steamboats began plying the Tensas River. A major population influx began in 1836, and the population kept growing until the Civil War, going from 5,142 in 1840 to 14,133 in 1860. The war led to a dramatic decline to a mere 8,600 in 1870. Over the next couple of decades, the numbers grew again to 14,135 in 1890. Changing economic conditions and the boll weevil outbreak caused another decline that was only reversed between 1920 and 1940, when the number of residents reached 14,826. By 2010, it had fallen to 12,093.

Cotton and the quality of the soil drove this influx. By 1850, there were 27 landowners in the parish who owned more than $20,000 (over $590,000 in 2015 dollars) worth of real estate. The largest holding was valued at $140,000 (well over $4,000,000 in today’s dollars).

According to Moncrief, “the newcomers cleared away the heavy forests and planted the new ground in the favored crop then, as now –– cotton. They cleared all the lands fronting water courses (which are the highest and most desirable lands for cultivation in this region) to form a continuous line of plantations along the streams.” Streams in this context refers not just to the Tensas but also to the smaller non-navigable bayous. Cotton raised along the smaller streams was brought down to the Tensas in flat-bottomed boats.

Moncrief’s thesis also includes figures for cotton and corn production in Madison Parish. Cotton production peaked at over 46,000 bales in 1858. (Pough was apparently incorrect in stating it was over 100,000 bales; he may have combined the total with that of an adjoining parish.) It had fallen to 1,830 by the end of the war. Production recovered between 1870 and 1875 and reached a postwar/pre-boll weevil peak of 25,981 bales in 1890, about the same level of production as in 1936. Corn production peaked at 618,620 bushels in 1859, falling dramatically after the war, peaking at 836,000 bushels in 1909, and then falling to 320,000 by 1936. My crude, back of the envelope estimate based on  yields of 5 bales per acre for cotton and 15 bushels per acre for corn, suggests that between a quarter and a third of the total acreage in the Parish was under commercial cultivation prior to the war.

While Moncrief’s paper evokes Tanner by describing the Singer Tract as 81,102 acres (126 square miles) of virgin timber, it also notes, “The tract includes several abandoned and grown up plantations, which after the Civil War, reverted to the state and were later sold to the present owners.” The ruins of one plantation house are still standing, deep within the Tensas National Wildlife Refuge.

While it was adjacent to and not strictly part of the Singer Tract, the story of the Frisby Plantation is illustrative. The plantation was established in the early 1850s with land acquisitions taking place over the next decade. Norman Frisby, the founder, was murdered by his nephew by marriage in an 1863 in a dispute over property. When Frisby’s widow was forced to sell the plantation in 1870, it totaled 19,479 acres, and its crops generated over $77,000 in revenue (the equivalent of $1.36 million in 2015). Tanner visited the site of the plantation and photographed one of its old fields. I haven’t been able to pinpoint the location of the old house; one 19th century survey survey seems to place it in Tensas Parish, near Fool’s River. Another account (from the history of the Sharkey Plantation discussed below) says it borders Disharoon (or Dishroom) Bend, much closer to the core of the Singer Tract. As shown on this overlay of 1875 land ownership on a modern map, the Frisby holdings included parcels adjacent to Mack’s Bayou and on Dishroom Bend.

The maps help flesh out the story. An earlier and incomplete parish-wide map of patentees shows that many parcels in the Singer Tract were purchased well before Frisby started acquiring land in the 1850s. Lands purchased in the 1840s include parcels along John’s and Mack’s Bayous, which makes sense since frontlands along streams were most desirable. This history of the Sharkey Plantation reveals that land sales began in the heart of Tanner’s search area during the 1840s. The author explains that the Sharkey plantation and others like it were more like communities, with a cluster of families (and presumably their slaves) living in close proximity, near a watercourse.  The 1875 map shows that much of the Singer Tract remained in private hands even after many parcels were abandoned during the Civil War.

Perhaps even more telling is this hand drawn map of Madison and Carroll Parishes from 1862. It shows the locations of towns, roads, ferry crossings, and plantations in the Singer Tract area. While it is incomplete and John’s Bayou is not shown, Sharkey Road is there, cutting in a southwesterly direction from the Richmond-Carthage road, crossing Alligator Bayou, and the Swearingen parcel. Another road crosses the heart of the Mack’s Bayou home range and the Tensas itself. The map delineates abandoned plantations and appears to show that, except for those abandoned areas, some cotton was being grown in every division of the Singer Tract. However limited the agricultural activity may have been in these sectors, the area was hardly a primeval wilderness; habitat had been fragmented; and old growth conditions were likely restricted for the most part to areas unsuitable for farming.

This passage from an 1885 article from the New Orleans Times-Picayune sheds some additional light on conditions in the area both before and after the Civil War. “But little has been said about Tensas River and Joe’s Bayou as, but little interest is there as compared with former years. Before the war there was a continuous planting interest all along those two streams but overflows and the war left them to grow up into weeds and bushes. In 1870 Mason, and later Loyd bought cattle from other parts of the country and carried them to those bayou places for pasturage, wherein a few years they made large sums of money. This was in the neighborhood of Quebec, which before the war was a flourishing little city, shipping 7090 bales of cotton. It was at the junction of the Tensas River and the railroad. It is now a waste place and to pass there on the railroad you would never know that a town had been there.” (In The Race to Save the Lord God Bird, p.76 Hoose plays into the virgin forest myth by claiming that railroads “finally reached the Tensas River sometime around 1900.”) Quebec was just a few miles outside the tract, near Bayou Despair, where Tanner listed a pair from 1934-1936.

The Rootsweb pages provide a couple of additional and important pieces of information.

Theodore Roosevelt visited Madison Parish in 1907.  Roosevelt’s descriptions provide added detail about conditions in and around the Singer Tract several decades after the Civil War. According to Roosevelt:

“Beyond the end of cultivation towers the great forest. Wherever the water stands in pools, and by the edges of the lakes and bayous, the giant cypress loom aloft, rivalled in size by some of the red gums and white oaks. In stature, in towering majesty, they are unsurpassed by any trees of our eastern forests; lordlier kings of the green-leaved world are not to be found until we reach the sequoias and redwoods of the Sierras. Among them grow many other trees–hackberry, thorn, honeylocust, tupelo, pecan, and ash. In the cypress sloughs the singular knees of the trees stand two or three feet above the black ooze. Palmettos grow thickly in places. The canebrakes stretch along the slight rises of ground, often extending for miles, forming one of the most striking and interesting features of the country. They choke out other growths, the feathery, graceful canes standing in ranks, tall, slender, serried, each but a few inches from his brother, and springing to a height of fifteen or twenty feet. They look like bamboos; they are well-nigh impenetrable to a man on horseback; even on foot they make difficult walking unless free use is made of the heavy bush-knife. It is impossible to see through them for more than fifteen or twenty paces, and often for not half that distance. Bears make their lairs in them, and they are the refuge for hunted things. Outside of them, in the swamp, bushes of many kinds grow thick among the tall trees, and vines and creepers climb the trunks and hang in trailing festoons from the branches. Here, likewise, the bush-knife is in constant play, as the skilled horsemen thread their way, often at a gallop, in and out among the great tree trunks, and through the dense, tangled, thorny undergrowth.”

The most salient point here is that Roosevelt’s “great forest” applied to low-lying areas in which there was standing water (something that Hoose glosses over). Roosevelt also saw three Ivory-billed Woodpeckers:

“The most notable birds and those which most interested me were the great ivory-billed woodpeckers. Of these I saw three, all of them in groves of giant cypress; their brilliant white bills contrasted finely with the black of their general plumage. They were noisy but wary, and they seemed to me to set off the wildness of the swamp as much as any of the beasts of the chase.”

 

A photograph from the hunt is here. Tanner seems to have been unaware of the Roosevelt encounter. Roosevelt’s visit came just 17 years into cotton farming’s second decline and 52 years after the end of the Civil War. Habitat conditions are likely to have been poorer in general than when Tanner was there 3 decades later. The relative ease with which Roosevelt saw three ivorybills (despite their wariness) suggests they were not uncommon in 1907 and calls Tanner’s assumptions (pp. 48-50) about fire, tree death, and population influxes between 1911 and 1930 into question.

There’s another gem in the Rootsweb pages. It’s not directly on topic, but it relates to Tanner’s later dogmatism. In arguing for extinction and dismissing post-Singer Tract reports, many of which involved birds being flushed from tree stumps or other locations near the ground, Tanner characterized this behavior as being characteristic of pileateds not ivorybills.

Rootsweb has a newspaper account of  T. Gilbert Pearson‘s  visit to the Singer Tract in 1932. Pearson (who was President of the Audubon Society at the time) was the first ornithologist to confirm the presence of ivorybills in the Tract. He saw, “The birds . . . feeding on stumps of rotting trees, the tops of which had been broken off. A favorite place for feeding is also on dead limbs at or near the tops of the very tall sweet gum trees found abundantly in this region.”

The evidence that relates directly to Tanner’s study area and its immediate environs suggests that claims about “virgin forest” and IBWO dependency on old-growth are based on flawed premises. The Singer Tract was no doubt a remarkable place, a huge area of contiguous and relatively undisturbed forest, but it’s clear that much of it was not old growth or “virgin”.

It’s more useful to think about what the Singer Tract is likely to have offered Ivory-billed Woodpeckers – some measure of seclusion, enough big trees for roosting and nesting, and an abundance of standing and fallen deadwood. The myth that the Ivory-billed Woodpecker required vast tracts of “virgin” forest may be emotionally compelling, but it’s not based on evidence; it’s time to put it to rest.

I’m looking forward to spending a week in the field starting just after Christmas.


Tanner and Population Density

I recently gave a talk to the Rockland County Audubon Society, and a member raised what I think is the strongest question about our evidence and about the persistence of the Ivory-billed Woodpecker in general. “How could the species have survived in such low numbers and at such low densities?”

In other posts, we’ve pointed to evidence that Tanner missed a population in Mississippi and was somewhat cavalier in his approach to evaluating potential habitat, disregarding advice Herbert Stoddard gave him in 1936, “The area where they (Ivory-billed Woodpeckers) may occur at present is simply tremendous, not restricted as many believe.”

A recent study on Magellanic Woodpeckers points to another factor that raises even more doubt about Tanner’s estimated population of 22 in 1939.  The study was conducted in an old growth Lenga forest in Patagonia. According to this study: “Our results show that Magellanic Woodpecker family groups require a minimum of 100 ha in old-growth forest habitat; thus, forest patches in less favourable forest conditions (e.g., younger, managed, fragmented, mixed forests) should probably be much larger to support a resident pair or family.”

The specific criteria that Tanner used for estimating the 1939 population at approximately 22 are unclear, but he assumed a maximum carrying capacity of six square miles per pair. The Wikipedia entry on the IBWO is even worse and is generally rife with bad information; the editors there expand Tanner’s six square miles to “9.7“.  Elsewhere, Tanner suggested a minimum home range of two and a half to three square miles. These numbers are somewhere between six and 16 times the minimum for a large southern congener that, like the ivorybill, lives in more temperate habitat than others in the genus.

Thus, there is a strong possibility that Tanner severely underestimated ivorybill populations in Florida. If he was so badly wrong about home range, he’s more likely to have missed populations in areas that he rejected for being suboptimal and not expansive enough.

Double the minimum acreage required by the magellanic in optimal habitat and apply that number to the ivorybill, and Sherburne, a large Louisiana WMA, could theoretically be home to just under 90 family groups. (We don’t think this is the case.) Even at 450 hectares per family group, the carrying capacity in Sherburne would be nearly 40. Such numbers are improbable in the extreme, but 9 or 10 family groups in an area that size would be very hard to detect.

With significantly smaller home range requirements, a substantially larger population in 1939, and a recognition that Singer Tract-like conditions are not a requirement (as Tanner himself made clear), various survival scenarios become considerably more plausible, assumptions about low densities become more questionable, and the quantity of potential habitat is far greater than anyone has imagined.

Edited to add: Although the study referenced above is more recent, Noel Snyder made the same basic argument about magellanics and other congeners in The Travails of Two Woodpeckers (2009). Snyder posits that hunting pressure, not habitat loss, was the primary cause of the ivorybill’s decline.

Snyder (who to the best of my knowledge has little hope for the ivorybill) does not fully address how Tanner’s assumptions might have affected his population estimates and habitat evaluations. He also doesn’t consider how taking Tanner at face value has influenced both search protocols and the “credibility” of post-Singer Tract reports. Nevertheless, he does hint at what I suspect is the key to the species’ survival: “With food supplies degraded, not eliminated, a reasonable possibility appears to exist that many ivory-bill populations in logged regions might still have found enough food to persist and might have endured at modest densities, had they been free of shooting pressure. The long persistence of the ivory-bill in one quite thoroughly logged region in Cuba supports this possibility . . .”

In my view, Snyder goes a little too far in downplaying specialization as a factor, even if Tanner overplayed it. It’s pretty clear – from range, habitat, and morphology – that ivorybills are more specialized than pileateds. But if the IBWO did persist after World War II and Snyder is right that hunting was a major factor in the species’  decline (even in the Singer Tract), there may be even more room for optimism, since hunting practices changed considerably in the post-war era.

 

 


Some Thoughts on The Ivorybill’s Decline

Jamie Hill, who has worked with the Cornell and Auburn teams, recently posted a Facebook link to a very interesting article from the September 2014 issue of Smithsonian. Ivory-billed Woodpecker aside, the piece is well worth reading, but for the purposes of this blog, the article got me thinking about reasons for the ivorybill’s decline and the possible role of the longleaf pine. These ideas are not entirely new or original with me; Lester Short went even further, suggesting that pine might have been the ivorybill’s primary habitat; Jerome Jackson devoted several pages of In Search of the Ivory-billed Woodpecker to pines, and Fangsheath of the ivorybill researchers forum has hinted at this too.

I was struck by just how congruent the historic range of the ivorybill is with the range of the longleaf pine (Pinus pilastrus). The overlap is not exact, and the pre-Columbian range of the ivorybill extended as far north as Ohio. Nonetheless, conditions in the Singer Tract were objectively quite different from what they were in many other parts of the historic range.

A recent blog post on the Tallahassee Democrat site reiterates the conventional wisdom about the species and the reasons for its decline. Author Budd Titlow writes: “Before the Civil War, when much of the southeastern U.S. was covered with vast tracts of primeval hardwood swampland, ivory-billed woodpeckers ranged from North Carolina south to Florida, west to Arkansas and Texas, and north into Oklahoma and Missouri. Then, after the Civil War, extensive logging of these old-growth swamps wiped out most of the ivory-billed’s habitat in one fell swoop.”

While there’s some truth to this history, it’s also a stereotype that’s based in large part on an imperfect reading of Tanner’s monograph and even more on Tanner’s dedication to protecting the Tract as the last remaining extensive old-growth stand in the southeast (although the Tract contained considerably less old growth than Tanner believed). Tanner’s efforts were admirable; the loss of countless acres of magnificent old-growth swamp forest was devastating environmentally and is unquestionably something to be mourned, but it seems unlikely that the destruction of these forests was the primary cause for the ivorybill’s decline.

The species was known to be disappearing by 1890 or even earlier, and Chester Reed’s 1906 Bird Guide to Land Birds East of the Rockies stated that the birds were restricted to isolated parts of Florida and possibly to “Indian Country” (Oklahoma). In The Travails of Two Woodpeckers, Noel Snyder, who attributes the decline primarily to hunting, points out that intensive logging of bottomland hardwoods began between 1890 and 1900. Logging of pine forests began considerably earlier, and these forests were severely fragmented, even before the Civil War. Snyder reads the early record (I think selectively) as indicating that ivorybills strongly preferred bottomland hardwoods and seldom used pines, in contrast to the Cuban ivorybill and the Imperial.

Jackson takes a different view, citing multiple references to the use of pines for feeding and nesting. Where Snyder reads Alexander Wilson’s early account as reflecting a preference for “swamps and bottomlands”, Jackson reads him as describing the preferred Carolina habitat as “a mosaic of baldcypress swamp and pine uplands, similar to the habitat in Florida”. Jackson goes on to suggest that, “It appears . . . that ivory-billed woodpeckers will inhabit both hardwood forests of river bottoms and pine forests of higher elevations, particularly old growth forests supporting healthy populations of beetles. They seemed to do best at the interface of these forest types, taking advantage of the resources of each.” (Emphasis added).

This meshes well with what Allen and Kellogg observed in Florida in 1924; the birds nested and roosted in cypress and were observed and photographed foraging in open pine forest. The Lambs’ limited observations in Cuba suggest something similar, a preference for roosting in pines but an equal division between pines and hardwoods for foraging.

Thus, it seems possible that the Singer Tract was actually suboptimal habitat for the ivorybill, since it contained no pine and little cypress. I’m also led to suspect that habitat fragmentation, rather than habitat loss may have been central to the decline of the ivorybill, with hunting as one of several other contributing factors. This fragmentation actually began well before the Civil War, but it accelerated with the post-war destruction of the longleaf pine forests, followed by the logging of the bottomlands. I’m personally convinced that the species beat the odds and survived, using one or both of the strategies discussed in this post. I wonder whether some of the modern search efforts have focused excessively on the bottomland hardwood model and not enough on areas where there’s an interface between forest types.


George and Nancy Lamb’s 1957 Cuban Ivorybill Study

I have been re-reading George Lamb’s 1957 report on the Cuban Ivory-billed Woodpecker. A number of items struck me as potentially significant for North American searchers, some for how they diverge from Tanner and others for their level of detail. Since this report is likely unfamiliar to many, I thought I’d do a quick post listing some of the more interesting observations

Lamb references a number of local sightings of “groups” of ivorybills, with one report to John Dennis that involved six birds. Notwithstanding, Lamb estimated the population density in Cuba to be much thinner than in the Singer Tract, at one pair per 12-25 square miles. He also pointed out that “ . . . the Cuban Ivory-bills are living for the most part in a cut-over pine forest where only small and deformed trees remain.”

The Cuban ivorybills fed on pines and hardwoods more or less equally, although most of the feeding sign was found on pines, due to the difficulty of searching for sign in the denser hardwood habitat. Roosts and nests were found exclusively in pines (one unused cavity was found in a hardwood), which is interesting in light of the fact that hardwoods were also available. Cavities were found at heights ranging from under 20 feet to nearly 60 feet. Cavities were higher in mature forest; Lamb suggested but did not conclude that the preference was for higher cavities and that the lower ones reflected an adaptation to cut-over conditions.

Lamb describes a female scaling bark: “At this point she was only about 25 feet away while she was feeding around the base of a small pine. She began ‘barking’ this tree about 30 inches from the ground and slowly worked up to the top.” Dennis too had observed birds scaling small pines. They found more scaling than excavation.

This apparent preference for pines, including small ones, may be significant, particularly since the hardwood areas were “relatively untouched”.

An estimated 17 birds were killed by humans over a ten year period, a huge number for such a small population. And it seems an open question whether the thinner population density noted by the Lambs was due to habitat quality, hunting pressure, or a combination of the two.

Regarding flight style: “. . .the flight of the Cuban Ivory-billed Woodpecker was always level and purposeful. They are strong fliers, capable of covering considerable distance in little time, as indeed they must to live successfully in cut-over woodlands. Although the Ivory-bill did not seem to undulate in its flight, the wing beats were not steady, having an almost imperceptible 2-3-2-3 rhythm.”

There’s no mention of double knocks, but calls are discussed. Lamb describes the sound as like the “note of a penny tin trumpet . . . short and usually repeated in a series of single-double-single beats, or it may begin with a double call: that is a high nasal “pent, pent-pent, pent”, or just “pent-pent”. On several occasions the female Ivory-bill most frequently observed made a few long and very loud calls, soon after leaving here roost tree in the early morning. The notes were of greater duration than normal and were repeated in a series of sixteen to twenty-two kients.”

Food for thought . . .


Habitat Conditions in the Singer Tract

Late last year, I wrote a post entitled “More Minutiae – Habitat Quality and Population Density in the Singer Tract”. I had to follow up with a couple of corrections and elaborations based on insights others shared with me. In the interest of providing more clarity and coherence, I thought I’d do a new piece combining the three posts and expanding on them a bit. I won’t delete the originals, but this one reflects what I think is a more accurate understanding of the material involved.

Mack's Bayou Ivorybill nest tree. Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

Mack’s Bayou nest tree. Courtesy of the Division of Rare and Manuscript Collections, Cornell University Library

The initial post was inspired by the image above and the caption describing it as the “Third ivorybills’ nest”, one I had looked at but not closely until last year. When I did examine it carefully, I was struck by how open the surrounding area seemed to be. Then I started going through archival photographs and scrutinizing them a little more closely.

In browsing through the Louisiana Digital Library’s collection of Singer Tract photographs, I came across an image that I had missed, one of the bridge over John’s Bayou taken in 1940. Tanner (p.32) includes an ivorybill sighting from this immediate vicinity, just northwest of the bridge. What I find interesting about the photograph is that the forest along the road appears to be fairly even-aged and does not have the characteristics typically associated with old growth. It is similar to what can be found in many parts of Louisiana today. An image from along Sharkey Road taken in 1937 shows similar characteristics, although another shot from 1939 (probably taken east of the bridge) shows more impressive looking habitat.

Richard Pough wrote a follow-up report to the Audubon Society on the Singer Tract in 1944. It’s a very interesting document that raises some questions about Tanner’s work. Pough explicitly accepted “Tanner’s premises as to the feeding habits and habitat preferences of the ivory-bill”, but he also noted “[n]othing in Mr. Tanner’s study indicates that Ivory-billed Woodpeckers show any preference or marked dependence on trees of great size such as one would find only in a virgin forest. He found them doing 14% of their feeding on trees 3”-12” in diameter and 64% on trees under 24” in diameter.”

Pough pointed out that “Maps of the area as of 1846 showed much of the Tensas River in plantations and many cleared fields back from the river on some of the interior ridges. This development continued for another 20 years until the Civil War, by which time Madison Parish was producing 110,000 bales of cotton a year. As the Parish has never produced over 30,000 bales since the Civil War, one gets some idea of how much land is now occupied by second growth forest of approximately 80 years age.”

Pough found only one ivorybill, a female. He assumed, relying on Tanner, that this was the only one left in the Tract. He may well have been mistaken, since Gus Willett, game warden in the Tract, wrote Tanner about seeing a pair in November 1948 (although the exact location is unclear). Correction, Willett did not write the letter, although the report reached Tanner, as discussed here.

The lone bird Pough saw was either the John’s Bayou female or one of its offspring. According to Pough, this bird was probably not feeding in virgin forest, and his report specifically suggests that Tanner might have been mistaken about the maturity of some of the habitat in the John’s Bayou area. In 1941, Tanner had written that the remaining John’s Bayou birds were roosting and feeding in “virgin” timber. Pough’s description of this area (and it seems to be the same patch) suggests it was likely cultivated pre-Civil War. It was devoid of big sweet gums, which Pough deemed to be the best indicator of old growth conditions, but had many dying Nuttall oaks 12-20 inches in diameter. Nonetheless, Pough relied on  Tanner’s premises to conclude that “only a relatively small portion of the total area of the Singer Tract supported a forest suitable as habitat for these birds.”

To return to the material in the earlier blog posts, much of the discussion focused on home ranges and the distribution of nest sites.

This map, drawn by Tanner after the 1935 expedition, lists three nests – designated as nests II, III, and IV –within a mile or two of each other and in the vicinity of John’s Bayou. Nest II is the famous “Elm Rock” nest. The map also shows a tree which is designated “Nest (?) III Squirrel” (also mentioned in Bales, “two miles to the south of the first nest” and fifty feet up p. 45). This was outside the area Tanner designated as prime in 1941 (p. 91); it is approximately a half-mile from the John’s Bayou bridge.

I’ve discussed this issue in depth with someone who’s very familiar with Tanner’s notes. I’m now persuaded Tanner concluded that nests III and IV from 1935 were not nests after all and that he assigned the birds involved to Titepaper (Nest III) and Bayou Despair (Nest IV). Nest IV is apparently one that Kuhn found but was unable to re-locate. Why Tanner changed his mind about it remains a mystery.

It’s very difficult to piece together this fragmentary information, and the monograph muddies the waters a bit by presenting the home ranges of the birds as being quite discreet, perhaps a good deal more than they were in fact. I suspect that Tanner decided the cavities were actually roosts, (although neither one is mentioned in the monograph). If so, they would have been well outside the home ranges Tanner identified and closer to the core of the John’s Bayou range than to the core of Titepaper or Bayou Despair.

Nests I and V were located near Mack’s Bayou. Nest I is dated May 14, ’34 on the map, and Nest V is dated May 10, ’35. The ’35 nest failed. It is the one referenced above. Allen and Kellogg described it as being 45’ feet up, in a pin oak snag, in a natural clearing, although it has been suggested that the snag may have been a remnant large tree in area that had been cleared prior to the Civil War. The nest designated Nest I and dated May 14, 1934 appears to be the one Tanner described on p. 81 of the monograph, “located within 100 yards of the second nest found in 1935”; however, in the monograph, he gave the date as May 13, 1933.

It’s worth pointing out that the Mack’s Bayou nests were in an area that Tanner designated as “best” for ivory bills (even if it’s not clear whether it was truly old growth). Nonetheless, nests failed in 1933 and 1935, and the adult birds had disappeared by 1938, apparently after producing one fledgling in 1936 or 1937. This was before the logging began.

My intention in writing those initial posts was to get a clearer handle on population densities and habitat requirements in the Singer Tract. In retrospect, I’m not sure that’s possible, since Tanner’s observations were almost entirely limited to one family of birds in a population that was dwindling for unknown reasons. At the very least, Tanner’s statement that 7 pairs of birds required 120 square miles of virgin forest in 1934 is based on an inflated estimate of the amount of old growth in the Tract and his minimum estimate of 6.25 square miles per pair also rests on that flawed premise.

Pough observed, “ . . . the ivorybill problem puzzles me exceedingly, and I do not feel that Tanner’s report begins to explain the reasons for the drastic decline in this species.” As the 2014-2015 search season approaches, I can only hope that the question of how the species persisted will puzzle people exceedingly in future years.