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.

Screen Shot 2018-09-23 at 8.41.28 AM

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.

Screen Shot 2017-12-22 at 3.42.31 PM

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

Screen Shot 2017-10-08 at 4.05.13 PM

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.

Screen Shot 2018-08-19 at 7.31.28 AM

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:

Screen Shot 2018-08-18 at 10.43.04 AM

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.

Screen Shot 2018-09-21 at 1.32.16 PM

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.


Bits and Pieces Part 4: Rethinking Range and “Suitable Habitat”

Part 1.

Part 2.

Part 3.

Bottom line upfront: In pre-contact and early colonial times and into the 19th century, Ivory-billed Woodpeckers inhabited a more varied and expansive range than most people realize. The range probably reached from the southern tip of Florida to central Ohio, with the 40th parallel as the approximate northern limit, westward to St. Louis and perhaps along the Missouri River. As discussed in the previous post, the western limit of the range appears to have been somewhere around the 96 or 97th Meridian, in the southwest, and probably somewhat farther east at the northwestern edge, if unproven historic reports from the Kansas City area are valid. Outside of coastal areas, distribution appears to have followed riparian corridors to its outer limits, and there are no records from elevations of over ~1000′.

A couple of maps may be useful for additional information about forest, bioregion, and habitat types and may help with visualization along with my map of records from unexpected habitats.

This is a new avenue of exploration for me, and I’m not personally familiar with many of the areas involved. I need to do additional research on areas within Tanner’s range map, along the coasts and in Florida, to be sure my characterizations are accurate. I’m leaving some of these locations on the master map for now (but removing them from the graphic below which includes extralimital and edge of range records). The master map is thus subject to change, so view it with the caveat that the notations may be incomplete or inaccurate. This post will address locations that are outside the confines of Tanner’s 1942 range map, although he accepted some of these records in 1989.

 

Screen Shot 2017-10-13 at 10.57.52 AM

As in the previous post, my analysis relies on records accompanied by reported physical evidence (even if that evidence has been lost) and to Native American sites where remains found are deemed unlikely to have been trade items. I relied on the US Fish and Wildlife Service’s Recovery Plan, Appendix E, Tanner, Jackson, and a few additional sources.

Before undertaking a whirlwind tour of the locations involved, I thought I should return to the mythology discussed in the previous post. I chose my words poorly in describing that mythology: “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).”

In fairness to Tanner, he didn’t characterize cypress as being abundant in the Tract. (It was not.) And his overall view was that cypress-tupelo swamps were not good ivorybill habitat. Audubon, with his romantic, indeed gothic, language is the father of the cypress myth.

I wish, kind reader, it were in my power to present to your mind’s eye the favourite resort of the Ivory-billed Woodpecker. Would that I could describe the extent of those deep morasses, overshadowed by millions of gigantic dark cypresses, spreading their sturdy moss-covered branches, as if to admonish intruding man to pause and reflect on the many difficulties which he must encounter, should he persist in venturing farther into their almost inaccessible recesses, extending for miles before him, where he should be interrupted by huge projecting branches, here and there the massy trunk of a fallen and decaying tree, and thousands of creeping and twining plants of numberless species! Would that I could represent to you the dangerous nature of the ground, its oozing, spongy, and miry disposition, although covered with a beautiful but treacherous carpeting, composed of the richest mosses, flags, and water-lilies, no sooner receiving the pressure of the foot than it yields and endangers the very life of the adventurer, whilst here and there, as he approaches an opening, that proves merely a lake of black muddy water, his ear is assailed by the dismal croaking of innumerable frogs, the hissing of serpents, or the bellowing of alligators! Would that I could give you an idea of the sultry pestiferous atmosphere that nearly suffocates the intruder during the meridian heat of our dogdays, in those gloomy and horrible swamps! But the attempt to picture these scenes would be vain. Nothing short of ocular demonstration can impress any adequate idea of them. 

Hasbrouck (1891) perpetuated the emphasis on cypress and very low-lying locations:

Screen Shot 2017-10-12 at 2.09.37 PM

Screen Shot 2017-10-12 at 2.10.17 PM

Hasbrouck’s explanation of the Missouri extralimital records is odd. The locations given are well outside bald cypress range.

That aside, the myth discussed in the previous post is a composite. The emphasis on cypress originates with Audubon and Hasbrouck, and the emphasis on large tracts, old growth, oaks, sweetgums, and tree size is largely drawn from Tanner.

Here are the extralimital records:

Reedy River, South Carolina: Nest with eggs reportedly collected in 1896 and later lost. The location is in the Piedmont, south of Greenville, at an elevation of approximately 900′. The Reedy flows into the Salad, which flows into the Congaree. This report is listed in the Recovery Plan because it was accepted by Sprunt as “definitive” but is considered highly questionable.

Etowah Mounds, Georgia: Pre-contact site, elevation approximately 700′. Presumably not trade goods “but requires further discussion” per the Recovery Plan. The Etowah River is at the northern end of the Alabama River watershed.

Between Martinsburg, West Virginia and Winchester, Virginia: Specimen reportedly collected by Wilson ca. 1810. Elevation at Winchester is 725′ and at Martinsburg is 453′. This would appear to be in the Potomac watershed.

Moundsville, West Virginia: Two lower mandibles found in a pre-contact (early Common Era, 0-200) midden. Potentially trade goods. Elevation 696′. The location is on the Ohio River.

Philo, Ohio: Near the Muskingum River. Tarsometatarsus found in pre-contact midden, dates from ca. 1100-1500. Elevation 735′.

Sciotto County, Ohio: Same as above. Elevation of the site is 1050′, but the nearby Sciotto River is lower. The site is about 10 miles from the Ohio, and the elevation at the confluence is 533′.

Ross County, Ohio: Same as above. Also on the Sciotto River but farther upstream. Elevation 863′

Near Troy, Ohio: Near the Miami River, March 1804 sight record by Gerard Hopkins, a Quaker envoy to the Miami and Pottwatomi. The report includes a description, “. . . resembling the red headed woodcock of Maryland, except that its head is black and its bill ivory. ” (Leese 2001). Omitted from the map because it doesn’t involve a specimen, but included here because it adds weight to the archaeological records.

Franklin County, Indiana: Report of a specimen, 1869, now lost. Elevation at least 490′. Probably along the Whitewater River, an Ohio tributary. There are a number of early reports from Indiana, but no other reported specimens and no archaeological records. In light of Audubon’s collection at Henderson, KY (just across the Ohio River) as well as the Ohio records, it seems likely that ivorybills were present in parts of Indiana into the 19th century.

Henderson, Kentucky: Female specimen collected by Audubon in July 1810 and used as a model for his first painting of the species. Elevation 400′. This record does not appear in the Recovery Plan or Jackson, and it seems to have been overlooked by researchers. Audubon’s own notation describes the location as “Red Banks”, on the Ohio River at the northernmost limit of the cypress-tupelo association, well upstream from the confluence with the Mississippi. Article by R. Haven Wiley in Kentucky Warbler, May 1970.

Screen Shot 2017-10-13 at 1.13.04 PM

Stanford, Kentucky: Pair reported, with one specimen collected by a Colonel Fleming, 1790. Record accepted by Tanner in 1989. Elevation 942′. The Dix River, which flows into the Kentucky and thence the Ohio, is nearby.

Cahokia Mounds, Illinois: East of the confluence of the Mississippi and the Missouri, 1500s or earlier, tarsometatarsus. Elevation 490′.

Forest Park, Missouri or Vicinity: West of the confluence of  the Mississippi and Missouri. Specimen collected 1896 and in the collection of the Colorado Museum of Natural History. Elevation approximately 500′.

I find it interesting that Ohio is the state with the most pre-contact records involving likely food items – 3 out of a total of 4-6.

With regard to Missouri and the Missouri River watershed, Hasbrouck accepted reports from Fayette and Kansas City, and given the overall picture, this does not seem implausible. In addition, there were persistent reports from the vicinity of Lake of the Ozarks, in the Missouri River watershed, until the end of the 1940s; Tanner received information about Missouri reports from local Audubon society officers but apparently disregarded it. There’s no way to assess the validity of these old, anecdotal claims and no evidence to support them, but given this perspective on the historic range, they may be somewhat less far-fetched than it seems at first glance.

I’m heading for Louisiana soon. More after I return – a trip report and probably two additional installments.