draft 7/10/23 then 7/15 7/17
Abstract. The formal literature on the Ivory-billed Woodpecker, Campephilus principalis, was reviewed and analyzed to establish an estimate of total population before anthropogenic effects for the species.
Exploring past population dynamics of Ivory-bills provides more information on natural trophic processes and niches that can be economically important to forestry and silviculture. The potential value of the species has been ignored which lessened, and lessens the management response by federal, state and local governmental departments.
Ivory-billed Woodpeckers were estimated to have a population of 57,696 animals 18,000 years ago (YA). Taking the above number of pairs, 28,848, potentially producing 2.11 fledglings per year per pair (Tanner 1942) gives 60,869 fledglings estimated in an assumed average year.
Total hypothetical maximum population after an average breeding season is estimated by combining adults and fledglings; 57,696 adults plus 60,869 fledglings, equals 118,565 Ivory-bills post breeding, 18,000 YA. The population of all species of birds drops post breeding due to predation disease, age, food resources etc.
Introduction
The formal Ivory-billed Woodpecker, Campephilus principalis literature concentrates on the specie's range, description, specimen records, interactions with other species including people, life history based mainly on the Singer Tract, LA and modern presentations of sightings and evidence. In an extensive literature review few details on the total population of the species from any date or time scale other than the 1930's (Tanner, 1942) was located. Minimal information can be found about the numbers of these large woodpeckers before anthropogenic effects began in the southeast United States.
The only detailed study of Ivory-billed Woodpeckers. (Tanner 1942) |
The Ivory-billed Woodpecker, if at carrying capacity, is a keystone species of significant ecological importance. Primary cavity nesters, such as woodpeckers, support community biodiversity and especially secondary cavity nesters. Picidae are an integral component of forest ecology.
The species past or possible economic importance is potentially substantial, but ignored; sustainable forestry was mostly unknown in prior centuries or deemed unimportant since forests were erroneously assumed to be impeding agriculture or a source of infinite firewood, fence posts, board feet, game animals, etc.
Ivory-bills with their unique and powerful bill and body morphology begin the pivotal "Tree to Soil" cycle ~ 24 months or more before any other Picidae can remove bark from a senescent tree exposing the cambium and heart wood. An intact Ivory-billed population can hypothetically increase long term productivity by up to 5% in some southeastern US forests as primary productivity, natural decomposition and silviculture regeneration rates are accelerated.
NBP's artist D. Tattoli draws Ivory-bills unique bill. Green lines indicate that the IBWO has its eyes positionally lower on head than the Pileated Woodpecker |
Ivory-billed Woodpeckers are relatively efficient within the Picidae at physically removing the hard bark covering of stressed or dead trees speeding up forest cycles, while converting wood boring insects into avian biomass. The ancestor species of the three northern clade species likley had
C. principalis has been noted to eat small beetle larvae (Dendroctonus spp.) prevalent under bark of recently stressed or dead pine trees. These beetles are considered one of the most important causes of economic loss in forestry (Ungerer 1999). Ivory-bills are a natural control on forest boring insects but the USDA, US Forest Service and the USFWS are confronted with fragmented, managed forest blocks and policies that immediately default to costly pesticides, or tree removal to address insect outbreaks. A balanced approach that included Ivory-billed management over the last 60 years may have resulted in a better integrated, efficient and effective result if coordinated governmental forestry and wildlife goals are properly merged. Department wide and interdepartmental MOAs united in a sustainable goal could have greatly lessened the large costs of pesticides and increased forest output while retaining biodiversity.
The most efficient ecosystems for converting solar energy into plant material and vertebrate biomass are the complex assemblage of biodiversity in East Africa.
"The combination of volcanic soils combined with the ecological impact of the migration results in one of the most productive ecosystems on earth, sustaining the largest number of ungulates and the highest concentration of large predators in the world." Source: UNESCO--Productivity of the SerengetiThe Serengeti produces a much greater biomass of plants and animals than anthropogenically managed open spaces. The assemblage of animals there accelerates basal rates of primary production and regenerative cycles. The Ivory-billed efficiently contributed to forest regeneration for possibly a million years. The Ivory-billed is a relatively large, understudied, forest bird; a keystone species that likely can increase economic output of some US forests if properly managed.
The US Department of the Interior, the US Fish and Wildlife Service, etc., have not properly valued the potential irreversible economic impact if the species is perpetually lost. Government entities have historically and recently underfunded the research, management for, and protection of, this valuable natural resource despite being mandated to preserve economic, recreational and natural resources for present and future American generations.
Exploring past population dynamics of Ivory-bills provides more information on natural forest processes that can be economically important to forestry and silviculture; Ivory-bills are an important resource but of course being critically endangered their beneficial impact is now miniscule.
Using the data and constructs detailed here an estimate is calculated for the total Ivory-billed population circa 18,000 years ago (18K YA) before many people (Pre-Clovis or Clovis) had arrived in the SE USA. By the 19th century there were certainly less birds than 18 K YA as anthropogenic effects were substantial; habitat destruction combined with increasing and more efficient direct taking of birds had and was accelerating.
Tanner (1942, p.31) said this about recent historical times:
None of the earlier accounts of the Ivory-billed Woodpecker contained accurate or definite statements as to the abundance of the bird. Judging from the interest that naturalists and collectors had in the Ivory-bill, and the accounts they wrote, it was never common. Most writers mentioned the Ivory-bill as being a rare bird, or an uncommon one, and some heightened the difficulties they had in securing specimens.
And:
As an example in North America, there have been estimates of the American Bison population number that are derived from descriptions and direct field observations related to the size of large herds. From such recollections and accounts, historians, anthropologists, and zoologists have estimated the original or pre-settlement population size for the species.Audubon usually described the big woodpecker as being quite rare.
Any estimate of a derivative of a dynamic complex ecosystem will have uncertainties. For example, it is not realistic to presume that before the Pre-Clovis and Clovis cultures reached the southeast United States that the Ivory-billed population was stable. Floods, droughts, large ice or log jams, beavers, insect outbreaks and fire regimes caused landscape scale ecological impacts pertinent to the Ivory-billed population demographics.
Riparian forest aerial, a small portion of the Ivory-bills possible habitat today; the remaining acres are greatly reduced and of lower quality and heterogeneity. |
Some animals are relatively simple to count |
For the Ivory-billed, historical observations inclusive of birds killed are used as base line data yielding animals per area. Tanner and others have examined these basic densities. Ecological interpretations on what was the likely total range of the species such as typical impacts of latitude, ocean currents, plains effect, etc., on climate and carrying capacity are utilized here.
On age demographics, including the ratio of non-breeding adults to breeding in the few historical metapopulations studied, there is little data. Conclusions on age structure, with veracity, could be made by examining A. T. Wayne's extensive field notes and papers which were not available online (personal communications, 10/2022). They can be accessed by a visit to the Charleston Museum of Natural History. Combining Wayne's info et al. with a coordinated examination of most or all skins, noting plumage details, age, molt, etc. would provide some information on the numbers of animals in respective age cohorts of the species.
Here this work rationalizes, or perhaps idealizes, the following unknowns: The pertinent field studies' birds, at Wacissa and California Swamps, which were counted as breeding adults, may not have all been adults. This research, roughly negates or nets out two drivers as following: the number of adult birds that were missed in the verbal population estimates of the time equals the number of birds that were actually fledglings, but counted as adults.
Undercounting actual numbers via field encounters and number of birds shot is very likely even in formal field studies since detection rates are needed (Buckland, et al., 2010). A. T. Wayne was collecting rather than studying the birds; this makes Tanner's research work important in the context of the subject of demographics.
Three Requirements
1) 18,000 YA, is ~ 2,500 years after the accepted date for the glacial maximum, assumed in this research to be an interglacial period with climate somewhat similar to the 19th century with annual and seasonally ice-free land mass available to the Ivory-billed approximately the same
The temporal point, 18,000 YA is assumed to have had a similar climate, ocean level and seasonally ice-free summer land mass available to the Ivory-billed as was in the 19th century. By choosing 18,000 YA we matched as closely as possible the forest conditions of the three most significant studies of Ivory-billed population density. In general, these three forested areas were considered to have late seral forests with old growth characteristics similar to the habitat of 18,000 YA. Tanner, 1942 used "primitive" to describe these areas.
Congaree National Park, NBP performed point surveys in over 30 locations |
Typical Point Survey Conditions |
Here are articles on some of NBP's field research:
SUMMARY NBP'S IBWO FIELD WORK
NBP IBWO STUDIES in CONGAREE NP
Obviously there were no surveys of any type done 18,000 YA; reliance on the various data gathered much later is temporally adjusted 180 centuries.
These maps or similar can be used to model improved granular population densities depending on variables such as forest type which each can be assigned, with ecological analysis, different population densities.
This map closely represents the amount of forest that may have existed 18,000 Years Ago |
USDA forest type information can improve population estimate for the Ivory-billed |
In this research the small number of studies together with anecdotal information strongly supported that C. principalis density had a north to south gradient which is to be expected for a species with nestlings that are obligate insectivores. This enforces known ecological drivers on population demographics of many species related to latitudinal, climate related clines (Hut, 2013) with actual observations of Ivory-billed population density. This supports the method of having two different bird densities based on the many biotic and abiotic characteristics latitudinal clines encompass, to more accurately model the Ivory-billed Woodpeckers' population.
Included are several million acres of breeding habitat outside of Tanner's generally accepted historical breeding range map as the target time here is 18,000 YA. The geographical breeding range here is also influenced by writings of P. Kalm (IBWO seasonal to SW New Jersey in the 1700s inferring MD for this analysis), J. J. Audubon (states to Maryland in the 1800s, inferring NC and VA), T. Jefferson (places the species in Virginia inferring VA and NC) and Hasbrouck (historical range into central coast of NC).
Various references indicate that even in 1600 the Ivory-bills range was larger than this depiction. |
MD 1 M
VA 2 M
NC 5 M
SC 10 M
GA 10 M
FL 40 M
AL 20 M
MS 15 M
LA 30 M
TX 30
AR 15 M
OK 10 M
MO 5 M
TN 5 M
IL 2 M
IN 1 M
KY 3 M
TOTAL 203 million acres
To estimate the carrying capacity for these two equally sized cohorts, north and south, we started with Tanner's summary (Tanner, 1942, pg 32) which is based on actual field work by himself and A. T. Wayne in the 19th and 20th centuries.
"This gives us three estimates of the abundance or density of Ivory-bills in primitive areas: in Louisiana, seven pairs in 120 square miles or one pair per seventeen square miles; in California swamp in northern Florida, about six pairs in sixty square miles or one pair per ten square miles; in Wacissa swamps in northern Florida, about twelve pairs in seventy-five square miles or one pair per six and a quarter square miles."
Two separate estimates of the Ivory-bills abundance or population density, one for the northern half of the total range acreage and one for the southern half are summated to obtain a total population number.
Conclusions
Tanner's and J.J. Kuhn's Singer Tract observations and density results is used for the N half of the Ivory-bills range. One pair per seventeen square miles is the estimated density for the N half of the IB's range. Each pair occupied 10,880 acres. The N half of the Ivory-bills range is modeled at 101,500,000 acres. 101,500,000 acres divided by 10,880 acres/pair = 9,329 pairs.
This equals 9,329 pairs x 2 = 18,658 birds
For the S half the two densities Tanner deduced from A. T. Wayne's field notes are averaged for the Ivory-bill from the two Florida areas; this results in one pair per 8.125 square miles.
This equals 19,519 pairs x 2 birds/pair = 39,038 birds
Total = 18,658 birds + 39,038 birds = 57,696 Ivory-billed Woodpeckers, estimated 18,000 years ago.
The Imperial Woodpecker (IMWO) has a maximum historical population estimate of 8,000 according to the article: Status and conservation of old-growth forests and endemic birds in the pine-oak zone of the Sierra Madre occidental Mexico (1996). This reference reports the total species range for the IMWO was 39,920 square miles and each individual needed 13 square Km. Imperial Woodpecker Population Estimate
Considering the two congeners and adjusting for respective total species range in square miles yields:
IBWO 317,187 square miles
IMWO 39,920 square miles
The range of the IBWO is 7.9 times that of the IMWO and the two species respective total population estimates are 57,696 vs 8,000. The IBWO has 7.2 times the population of the IMWO in 7.9 times the area. This infers that carrying capacity of the two species per area is somewhat similar with the Ivory-billed needing about 10% more area per pair.
The world's largest woodpecker is the likely extinct Imperial Woodpecker, which averaged 58 cm (23 inches) in length and is estimated to weigh over 600 g (1.3 lb). The Ivory-billed Woodpecker is smaller, with a length of 50 cm (20 inches) and a weight of 500 g (1.1 lb).
NBP during modern Ivory-billed studies employing advanced attraction techniques, accumulating data over 650 points from many locations. In a few forests, and from year to year, these locations supported the pair range estimate in this paper; NBP found that a pair of Ivory-bills could have a range of ~ 10,000 acres.
Post-breeding population estimate for Ivory-billed Woodpecker:
Obviously total population of all species of birds is maximal right after the breeding season. Populations can literally double or more in a few months; the increase is due to eggs hatching and fledglings leaving the nests.
At the Singer Tract, Tanner and Kuhn reported observing 6 Ivory-billed nests which had 9 broods from 1931-39 (Tanner, 1942 p. 81). The Ivory-bill had 19 young leave the nest from 9 broods; this is 2.11 fledglings/brood. Three broods failed. This is direct Ivory-billed field data on success of their breeding phenology. This is a small location sample size (n = 1) however the data was accumulated over several years with several nesting pairs.
Taking the above total pair number for the SE USA 18,000 YA we have 28,848 pairs potentially producing 2.11 fledglings per year per pair. This equals 60,869 fledglings.
Total population after breeding season is estimated at 57,696 adults plus 60,869 fledglings which equals 118,565 Ivory-bills.
On average these young birds will have a high mortality rate in their first year, while the adults will have a lower mortality rate. After the birds are fledged the total Ivory-bill population would have dropped until the next years breeding season due to natural mortality.
Ivory-billed nestling about 8 days before fledging |
Discussion
Objectively, from the totality of the C. principalis reports, more southerly populations within the range were specifically highlighted for their relative abundance compared to all other areas (Tanner, 1942). This enforces known ecological drivers on population demographics of many species related to latitudinal, climate related clines (Hut, 2013) with actual observations of Ivory-billed population density. This supports the method of having two different bird densities based on the many biotic and abiotic characteristics latitudinal clines encompass, to more accurately model the Ivory-billed Woodpeckers' population.
Two separate estimates of the Ivory-bills abundance or population density, one for the northern half of the IBs range and one for the southern half of the total range are summated to obtain a total population number in those acres.
Tanner, used his own observations and others (J.J .Audubon, A. T. Wayne, et al.) to relate that the peninsular Florida, Gulf regions of the Suwanee River such as California Swamp, Wacissa wetlands, and Buffalo Bayou (present Houston. Tx area) had a relative abundance of Ivory-bills. These three areas are all in the southern half of the Ivory-billed's range as defined in this research:
Location --------------Latitude (S to N)
Buffalo Bayou-------29.7400373,-95.357490
California Swamp---29.7612452,-83.054545XXXX
Wacissa Swamps---30.237583,--83.984288
Singer Tract--------- 32.3016241,-91.9054949
Tanner and his Cornell professors overestimated the amount of virgin forest in the Singer Tract; this infers the same forest, 18,000 years ago, may have supported more IBs than J. J Kuhn's Singer. For the N half of the IB's range the winters are colder and trophic resources available would have been reduced and less varied comparatively to the S half of the range. Intraspecific carrying capacity proportionally drops for more northerly latitudes for many semi-tropical and temperate taxa.
The two prior inputs or factors, one, underestimating the maximum carrying capacity of the Singer Tract by Cornell's/Tanner and, two, the decreasing carrying capacity with more N latitudes, are assumed to offset each other's carrying capacity impact for the N half of the IB's range. Further research into the differences in carrying capacity in relationship to latitude in a semi-tropical zone of North America will likely result in a better estimate of carrying capacity and population but any adjustments to the final estimated numbers should be minimal.
Ivory-billed Woodpecker in cavity. Copyright to Academy of Natural Science VIREO |
Boyer W. D. (1990). Pinus palustris Mill. Longleaf Pine, Pinaceae -- Pine family Boyer
Hut R. A. (2013)
Latitudinal clines: an evolutionary view on biological rhythms
https://royalsocietypublishing.org › doi › rspb.2013.0433
Lammertink. Status and conservation of old-growth forests and endemic birds in the pine-oak zone of the Sierra Madre occidental Mexico (1996). This reference reports the total species range for the IMWO was 39,920 square miles and each individual needed 13 square Km. Imperial Woodpecker Population Estimate
by · Cited by 239 — We review literature with respect to latitudinal clines
end of virrazzi paper
Raw Data
Beavers---to Fv trapping significant before 1800
Probably because of their feeding and breeding dependence on large old trees, great slaty woodpeckers are most common in primary forests and show density reductions of over 80% in logged forests.[6] The global population is in decline because of the loss of forest cover and logging of old-growth forest throughout its range, with habitat loss being particularly rapid in Myanmar, Cambodia and Indonesia which are the countries that still hold the majority of the global population. In 2010, the great slaty woodpecker was included in the IUCN Red List in the Vulnerable category.[1]
We conclude, first, that spatial scale has indeed had an important effect on the characterization of the Panama bird community. The intrinsically patchy distributions of most forest-dwelling bird species raise the need for large-scale censuses. Second, the Panama community, compared with the two Amazonian sites, has a fundamentally different organization; it hosts nearly twice as many individual birds and is distinctly less dominated by rarity. Similar patterns of community structure appear to be present within tree and mammal communities as well. Therefore, results from the Amazonian studies cannot be generalized to all lowland Neotropical communities. We attribute differences in community structure primarily to differing biogeographic histories. The lower species richness and the greater number of total birds present in Panama appear to derive, at least in part, from two important factors: an area effect linked to the location of Panama on a narrow isthmus, and the repeated history of disturbance on multiple temporal scales in Panama.
FB 9/12/22
PIWO may have 150 times the popalation of IBWO at one time; in the SE USA the ratio would been much less elevated.
Behaviorally the modern IBWO is not the same as the precontact animal. Due to nominal microevolution caused by severe and persistent anthropogenic taking of the least wary bird (subsistence, museum, and curiosity hunting) the bird now instinctively is "hard-wired" to prefer the most secluded section of forest which may be far from quality feeding areas for themselves or nestlings. Today the habitat landscape is dominated by short to longer, narrow river forests. The IBWO being instinctively fearful of people results in it foraging away from habitat patch/forest edge as you would if you instinctively were programmed to avoid, for example lions, which are in the fields and shade of the forest edge and they will kill you. So the aerial of a 10 mile long and 1 mile wide river forest habitat may seem to be 10 sq miles but an IBWO may perceive it as only 5 sq miles further reducing carrying capacity.
Finally the IB was already showing a loss of genetic variability before 1900. It's likely that there is some increase in embryological or hatchling mortality via various genetically based mechanisms that have increased the chances of homogygosity of deleterious alleles.
Regardless ecology is complex; the IBWO has somehow persited, microevolution is rapid and things can by definition change rapidly for the better.