Historic Range Hypothesis: A Bits ‘n’ Pieces Epilogue

In retrospect, I realize that I could have stated my hypothesis about historic range more explicitly in the Bits ‘n’ Pieces series, here, here, here, here, here, and here. My treatment of this subject draws on and expands upon Tanner, Jerome Jackson’s extensive review of the historical record, the work of Benjamin Leese, and Appendix E in the U.S. Fish and Wildlife Service’s Ivory-billed Woodpecker Recovery Plan, supplemented by my own research and review of the source material.

I got somewhat too focused on specimen records and even more so on habitat types. The version of the map I created for the series reflects that focus, by including barrier islands, for example. The subjects of range and habitat are of course intertwined, but I’ll state the hypothesis with respect to range upfront and follow it with a more detailed discussion:

The Ivory-billed Woodpecker’s range was considerably more extensive and than is recognized by the general public and than has been represented in much of the literature, both popular and scientific. All published range maps of which I’m aware, including Hasbrouck’s (1891), Tanner’s (1942), Jackson’s revised and redrawn version of Tanner (2004), and online, poorly reflect the historic range, especially east of the Mississippi.

This has implications in terms of habitat requirements and adaptability as well.

I suggest that the northern limit of the ivorybill’s range was just above 40 degrees North and that the western limit was approximately 98 degrees West. There are no archaeological or other records from over 2000′, so I’d exclude higher elevations in the Appalachians and Ozarks. I think the previous maps are accurate to the extent that they show the range as extending farther west in the Red and Arkansas River basins than in the Missouri.

To express it somewhat differently, I think ivorybills could be found as far north as the lower Delaware River on the Eastern Seaboard and that they could be found in riparian corridors into the lower reaches of the Appalachians from there south.

In Florida, I think it’s conceivable that the range extended to the Upper Keys. In the southwest, I’d draw the line near Port Lavaca, Texas to west of Tulsa, Oklahoma. Except for an archaeological find in northeastern Nebraska that almost certainly involved trade items, the northernmost specimen record from west of the Mississippi is from Forest Park, MO (more on that below).

The story seems to have been somewhat different in the Ohio River watershed, where both archaeological evidence and sight records suggest that birds were present far upstream from the Mississippi confluence and along tributaries well north of the river itself.

I’ve created another map showing both the limits of my hypothesized range and selected reports from what could be considered the edges of that range. A screen cap is below. Click on this link for annotations. Although some of the records are pre-Columbian (the earliest estimated as dating to 0-200 CE/AD), I suspect this was the approximate range until the mid to late-18th century.

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Hypothesized limits of Ivory-billed Woodpecker range and selected records from areas outside those shown on most other maps.

Thus my hypothesized range is considerably more extensive than the conventional one (shown immediately below). On the Eastern Seaboard, it extends much farther north and also farther inland. It is far greater within the Mississippi watershed, extending to north of Columbus, Ohio, and encompassing all but the higher elevations in the Appalachians (based on the archaeological site at 2000′ near Lebanon, VA), reaching farther north and west in Arkansas and Missouri.

It was also somewhat more extensive in Texas and Oklahoma, reaching farther into the plains and a little farther south than the maps suggest. Outside of coastal areas, the presence was probably limited to riparian corridors, with those in the western part of the range (at least) being narrow. The version below is from the IUCN Red List entry for the ivorybill.

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IUCN Map derived from Jackson’s revision of Tanner’s 1942 publication. Similar maps are widely reproduced.

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Hasbrouck’s Ivory-billed Woodpecker Range Map (1890). Shaded areas reflect what Hasbrouck believed to be the range at the time of writing.

While what I’m suggesting may seem heretical to some, it’s well-supported by the archaeological record, accounts from early explorers and naturalists, and collection records or reports thereof, albeit to a lesser extent. I covered many of the records that led to my formulating this hypothesis in previous posts; I won’t recapitulate all of them here, but one passage is worth revisiting. Two notes on technical terminology: middens are most easily defined as “dumps for old domestic waste“, and tarsometatarsi are the lower leg bones found in birds (and some dinosuar fossils).

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. 

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 uncovered 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.

In an unpublished 1989 update to his monograph (housed in the archives at Cornell), Tanner accepted additional reports, including archaeological finds from Scioto County, Ohio (15th or 16th-century) and Madison County, Illinois (Cahokia Mounds, approximately 15th-century), as well as a collection record from Forest Park, Missouri and an 18th century record from Lincoln County, Kentucky. Since that time, additional archaeological records from Georgia, Virginia, West Virginia, and Ohio have turned up.

Thus, the case for ivorybill presence both in the lower reaches of the Appalachians and well up the Ohio River (and its tributaries) is compelling, and any suggestion that items found in Native American middens might be trade goods is pure speculation with no evidence to support it.

In a paper titled, “Putative Records of the Ivory Billed Woodpecker in Pennsylvania and the Mid-Atlantic” (2016 Pennsylvania Birds, 20(2):71-72), Leese suggested that eastern Pennsylvania and Southern New Jersey might be the northern limit on the Atlantic Coast. I agree and might even go a step further and argue that the available evidence strongly supports that view. The discussion below relies heavily on material Leese discovered, though I disagree with Leese’s conclusion that a more extensive range and more varied habitats support Snyder’s argument that hunting was the “main cause of the species’ extinction”.

This idea that ivorybills ranged so far north on the East Coast will probably be the hardest for many to swallow. Bear in mind, however, that Audubon described the ivorybill as an accidental in Maryland. Audubon’s discussion of range is odd; he didn’t mention Florida, and there’s nothing to support the idea that the western part of the range extended to the foothills of the Rockies. But he would have been far better informed about the Eastern Seaboard.

In any event, his description dates to the 1820s, by which time the range was likely shrinking. As discussed previously in the series, I suspect that the extirpation of the beaver played a major role in the ivorybill’s decline outside of Florida. Beavers were probably extirpated in New Jersey by 1820 (Van Gelder 1984). It’s reasonable to infer a similar date for eastern Pennsylvania.

To expand on this beyond the previous posts in the series, Peter Kalm, a prominent European naturalist and student of Linnaeus, listed the ivorybill as present in the Delaware Valley and distinguished it from the Pileated, which he described as “a Black Woodpecker with a red head” and “frequent in the Pennsylvanian forests”.

It’s not clear whether Tanner was aware of Kalm’s 1749 record, which was described in The Auk in 1903. The author of that paper suggested that this record should be taken “cum grano salis,” primarily because there have been no other records from the state. Or have there?

While it is ambiguous, Leese discovered a letter from Alexander Wilson, written during the early years of his career, when he was living on William Bartram’s property near Philadelphia; it is strongly suggestive of ivorybill, describing a large and “most extraordinary Blackheaded Woodpecker”. This was at a time when Wilson seemed to be in the process of learning his birds, and he did not mention it in later correspondence, which leaves room for doubt.

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While Wilson’s description is not very detailed, I find it suggestive. And as noted previously, I think the only basis for rejecting Kalm’s account (and he seems to have otherwise been meticulous) is what we think we know about the ivorybill, a knowledge base derived from observations dating to between 60 and 190 years later. While there’s no way to prove it, I think Kalm’s listing should be treated as credible; it would be if a more common species were involved. Leese mentioned a couple of additional 18th century reports from Pennsylvania (included on the map) but found them questionable.

I’m not aware of anyone else having made this observation, but I think it’s a very important one. In the 1740s, the ivorybill had not yet acquired the mystique that would accompany it from the mid-1800s to the present, a mystique that was popularized if not invented by Audubon. This lends greater credibility to earlier reports such as Kalm’s. I think the same principle supports Jefferson’s listing of the ivorybill as resident in Virginia (which may have been based on his own observations) a couple of decades after Kalm and a few decades before Audubon.

To return briefly to Wilson and another report from an unexpected location, Jackson (2004) references a specimen collected by Wilson from somewhere between Winchester, Virginia and Martinsburg in what is now West Virginia (presumably in the Potomac watershed). I have not located the primary source for this reference, which may be in the archives of the Philadelphia Academy of Natural Sciences, but here’s Jackson’s:

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The Putnam County, WV find is also interesting, as it suggests a bird that may have been collected for trade rather than being acquired through it.

I want to touch briefly on two additional records, both involving specimens. The facts related to these records point to how even specimen evidence is not free from ambiguity. For one thing, specimen tags often reflect shipment rather than collection locations. In addition, labels could have been changed and replaced at any point in the chain of custody between hunter and museum.

The first specimen is housed at Cornell (and I have seen it myself). It was collected in 1898 from the “Florida keys”. In Appendix E of the Ivory-billed Woodpecker Recovery Plan, it is suggested that this might actually refer to one of the forested “keys” in the Everglades rather than to one of the offshore islands. However, given the ivorybill’s use of barrier islands on the Atlantic Coast and the 10,000 Islands in Florida, it does not seem farfetched that the specimen could have been collected somewhere in the Upper Keys proper.

From my current perspective, based on the Recovery Plan’s information about habitat on the southwest Florida coast, both possibilities seem plausible and neither is particularly surprising. However, neither location would ever have had many characteristics in common with stereotypical ivorybill habitat –  vast, contiguous bottomland hardwood forests. (Although he sometimes seemed to embrace this stereotype later in life, Tanner recognized that ivorybills lived in varied habitats, as have others who have studied the subject in depth.)

The other specimen, labeled  as being from Forest Park, Saint Louis, Missouri, escaped Tanner’s notice initially but was included in his 1989 update. It is housed at the Colorado Museum of Natural History in Denver and was collected in 1886. Matt Courtman pointed out that by 1886, Forest Park was hardly an isolated spot; it attracted “hundreds of thousands” of visitors a year,was located on the outskirts of a city with a population of 400,000, had been at least partially landscaped, and was accessible by streetcar.

So not only did this collection come from well north of what’s commonly accepted as the historic range; it may have come from a managed parkland on the outskirts of a major city, hardly stereotypical ivorybill habitat. The location and date are surprising, so it’s possible that the specimen was collected somewhere else (though not in a different region) and that Forest Park was the shipping location.

This may be so, but an 1886 collection from anywhere near Saint Louis is a paradigm-buster, regardless of the exact location, as this 1884 map suggests. Forest Park, to the southeast of Florisant and Ferguson, which are indicated by the red circle on the map, is not shown.

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Detail from 1884 map of Missouri, showing Saint Louis and environs. Note both the network of railroad lines and the apparent population density.

Tanner’s original map showed the ivorybill as never having ranged farther north than the Missouri bootheel (the southeastern corner of the state) and as having been extirpated from all but the southernmost tip of that region by 1880. The last specimen collected in Missouri dates to 1895, from just north of the bootheel “eight miles southwest of Morley.” (Widdman, 1908). Tanner seems to have been unaware of this record, which is mentioned in Jackson, who could not locate the specimen itself.

Tanner received at least one report (1937) from Dallas County, Missouri, which is northwest of Springfield, in the upper (southern) reaches of the Missouri River basin and well outside the generally accepted historic range. The source was a local Audubon Society officer, and the letter is archived at Cornell. There appears to have been no follow-up. According to the Recovery Plan, reports from this general area continued until 1949.

Though his map of the range ca. 1890 suggests a northernmost limit in Mississippi and Louisiana, Hasbrouck (citing Cooke, Bird Migration in the Mississippi Valley) mentioned 1884 records from near Kansas City and undated claims of former breeding near Fayette, along the river in central Missouri; Tanner considered these questionable or accidental. I agree that they’re somewhat questionable but not that they could have been accidental, since both purportedly involved breeding.

Reading Cooke reveals that both were second-hand accounts. The Kansas City report includes no details except that birds were observed “during the past few winters”, and “it probably still breeds in that vicinity”. For Fayette, the source was identified as a local farmer and egg collector by the name of Lientz, but the former breeding claim is devoid of any details, including the informant’s name.

Thus, the possibility of mistake or miscommunication exists in both instances, as in the seemingly credible report from eastern Nebraska discussed in Part 2; however, Jackson also references Harris’s Birds of the Kansas City Region (1919), which lists a “Judge Guinot” and others” for the Kansas City area records, in the “deep woods of the Missouri bottoms”. No date is given, but it seems possible if not likely that Guinot (1855-1935) was Cooke’s source

While I think the historic range probably extended as far as Kansas City, the case seems a little weaker to me than it does for the eastern seaboard and Ohio Valley –  given the paucity or absence of archaeological evidence or early reports from the Missouri River watershed, a key route for explorers, traders, and early settlers. For example, Featherstonhaugh, who crossed the Missouri River basin near the confluence with the Mississippi in the 1830s, described the northern limit as being considerably farther south, in the Arkansas-White River watershed. But perhaps there’s more to uncover about the ivorybill’s history along the Missouri.

As with anything ivorybill related, it’s probably best to be comfortable with not knowing and even to revel in the uncertainty.

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The Choctawhatchee: A Detour

I’m still planning a post on historic range and one on questions of evidence but thought I’d take this brief detour first. Tommy Michot is braving the Louisiana summer to change batteries and cards and to deploy an additional trail cam. We’re trying to service the cameras and replace the cards on a bimonthly basis. If there’s anything noteworthy on the cards, I’ll adjust my posting schedule accordingly. Look for the historic range post within a couple of weeks and the one on evidence a few weeks after that, before the start of search season in October.

This is the first time I’ve devoted an entire post to someone else’s effort. Though I’ve received a number of other intriguing reports, I’ve chosen to write about this one for a couple of reasons. First, I want to counteract the oft-repeated notion that reports have dried up in areas where intensive searches have taken place. Second, the searcher in question has found intriguing feeding sign as recently as 2017. The images included in this post are among the most suggestive I’ve seen from outside our search areas and tick most of my Ivory-billed Woodpecker boxes. I use the word ‘among’ advisedly here, since virtually all the feeding sign imagery that I’ve found intriguing comes from the Choctawhatchee, including the images showing extensive work on this page from the site devoted to the Auburn search.

The source of the report is an experienced birder and photographer named Rick Sellers. He has generously agreed to my posting this and allowed me to include some of his photographs. His first sighting was in 2012, approximately four years after organized searching came to end in the area and approximately seven miles downstream from Auburn’s ‘hot zone’. Rick shared the details with his family members and with Geoff Hill at the time and posted his email to his family on ibwo.net in March, 2014:

2/26/12
No doubt about it! While in the swamp today, I heard a large woodpecker hammering in the direction of a stand of slash pines at the edge of the swamp. I headed that way and just as I entered the clearing I saw the silhouette of a large dark bird leaving a tall tree on the other side of the stand of pines. I couldn’t ID it because the overcast sky was too bright. All I saw was the dark silhouette against the sky but the bird was clearly larger than a pileated woodpecker and flew loon-like, not undulating like a pileated. There had been a fire in recent years in this area and about 50% of the pines were dead showing extensive bark scaling, diagnostic of ivorybill foraging. Lamenting the fact that I had been unable to ID the big bird, I decided to stake out the pine stand, hoping that an ivorybill would return to feed. I found a secluded spot on the edge of the pines next to the swamp, sat down and ate lunch. I sat there for about 45 minutes and then as I was feeling rather drowsy, lay back with my head on my daypack. I was about to doze off when I heard, “kent-kent” coming from the swamp to my right, no more than 100 feet away! Thinking I must have been dreaming, I sat up and listened intently. Then I heard it again, “kent-kent-kent…..kent-kent! Over the next 3 or 4 minutes, I clearly heard 15-20 kents, some louder than others, that sounded exactly like Dan Mennill’s recordings from 2007. There was no doubt in my mind that I was hearing at least one, if not two, Ivory-billed Woodpeckers! Then as suddenly as they started, the kents stopped. I waited about 5 minutes before moving and then walked into the swamp in the direction of the kents. Unfortunately, I neither saw nor heard anything more as I walked around the area from whence the kents came. I plan to go back tomorrow bright and early and stake out the pines again. (end of email)

Since that encounter, I have been back to the Choctawhatchee at least 10 times for a week at a time. I have hiked and kayaked many miles but have had no more encounters. I am not discouraged, though. Just that one encounter is enough to keep me going until I can get the video or photo of the bird.

Rick informs me that he has had one possible sighting since the time he posted – a large woodpecker showing a lot of white – but that his confidence level is only around 50%.

Rick suspects that the birds do much of their feeding in upland pines outside the floodplain, which is where he had his sighting. He shared a couple of images from the location of his sighting with the notations showing the bird’s path.

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Some of the scaling in this stand of pines is extensive, but none of it strikes me as being beyond the capacity of a Pileated Woodpecker. On its own, the work shown in these images would be unlikely to pique my interest. But as in our search area, fire killed pines in surrounding uplands are, at least potentially, a major food source.

What really captured my attention were a couple of photographs. I found the first on Rick’s Facebook page. It was taken in 2017. The tree may be a tupelo, but I’m not sure. The bark is thin, and regardless of species, it is undoubtedly weaker and more easily scaled than hickory. There are also some hints of layered removal, akin to blonding. Nevertheless, a number of characteristics suggest Ivory-billed Woodpecker to me – the mostly clean edges, the lack of damage to or excavation of the underlying sapwood, and the targeted expansion of already large exit tunnels. This is unusual work, and it’s what inspired me to reach out to Rick for more information.

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Rick subsequently shared images of a scaled spruce pine he found in 2017. And while I’ve generally taken the view that there’s no way to recognize Ivory-billed Woodpecker work on conifers, this sign is strikingly similar to the work on hickories that we’re finding and also to the work of other Campephilus woodpeckers. The work is very extensive; there’s virtually no blonding or damage to the sapwood, except for targeted digging around the exit tunnels. It ticks my ivorybill boxes, save for the fact that it’s on a softwood and there was no chance to examine the bark chips. The final image below is a detail from one of our hickories for comparison.

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Except for a passing claim on Facebook about recent ivorybill sightings along the Pea River (a tributary) in Alabama, I’m not aware of other reports of sightings or auditory encounters in the area, but the fact that Rick has continued to find suggestive feeding sign, as recently as last year, suggests to me that the Choctawhatchee merits more attention than it has gotten since Auburn left. Of course, the same is true of many other areas, but this is the only instance where I’ve seen feeding sign that I strongly suspect is the work of Ivory-billed Woodpeckers. If I were looking for a place to search, the Choctawhatchee would be at or near the top of my list.


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; Matt Courtman blamed this tendency on internalized beliefs. And for my part, I sometimes find it hard to shake these beliefs.

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.

Back in October, Matt Courtman discovered that 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.


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.