The Sacred Sacred Ibis [reposted]

BY: Bob Montgomerie, Queen’s University | 15 October 2018

Thoth and ibises

The ancient Greeks usually depicted Thoth—their god of writing, wisdom and magic—as having the head of a bird with a long, down-curving bill.  Until the 1800s, Europeans thought that this bird was probably a curlew, a stork or a heron. Linnaeus believed that the bird must be the Cattle Egret which he called Ardea ibis in the 1758 edition of his Systema Naturae. It was not until the turn of the 19th century that a small group of French scientists and naturalists finally confirmed the connection between Thoth and the head of the African Sacred Ibis (Threskiornis aethiopicus). This species was not unambiguously described until 1790 [1], but it took Napoleon Bonaparte’s military campaign to Egypt to provide the connection between this species and Thoth.

We now know that the sacred ibis was considered (and revered) by the Egyptians to be the earthly form of Thoth. For about a millennium starting in 1100 BC, ibises were frequently mummified as an offering to Thoth, believing that mummification would put the birds on a direct line to the afterlife. As a result, several million sacred ibises were killed, gutted, embalmed and folded with the bill tucked between the tail feathers. The carcasses were then wrapped with linen dipped in resin, and inserted individually or in pairs into urns that were placed in vast underground caverns in cities all along the Nile. Many of these mummified ibises have grains, snakes, snails and other foods in their body cavities, possibly to provide the birds with some food in the afterlife.

But why ibises, and where did all of these birds come from? There can be no doubt that the sacred ibis was a reasonably common bird [2] in swampy areas all along the Nile in the Late and Ptolomeic Periods of ancient Egyptian civilization [3]. Those birds were of great value to nearby villages as they ate the snails that infested fish ponds, snails that harboured parasites dangerous to humans. They were also claimed to feed on flying snakes (?) and generally consumed all kinds of human refuse [4]. No wonder they were considered to be sacred.

At several sites of ancient cities along the Nile, archaeologists have found incredible numbers of mummified ibises: 1.75 million at Saqqara, 4 million at Tuna el-Gebel, for example. Even over a period of 500 years that is a lot of birds per year, likely magnitudes more than could have been hunted in the local marshes for any sustained period. Because of their religious importance, sanctuaries dedicated to the ibis sprang up all over the country, where birds were bred and raised in captivity, processing as many as 20,000 ibises per year for the votive ibis industry. Priests apparently gathered eggs for artificial incubation and tended the large flocks, as well as engaging in a large pottery industry to make urns for the mummified birds. These ibiotropheia may well be the earliest examples of bird-farming that did not involve some form of fowl.

The vast stores of ibis mummies in Egypt were brought to light by Geoffery Saint-Hilaire and Jules-César Savigny, two of the 167 savants [5] who accompanied Napoleon Bonaparte to Egypt from 1798-1801. Savigny noticed that the ibis often appeared in hieroglyphics and tomb paintings, and reasoned that this bird was important to Egyptian culture. He wrote up his discoveries in 1805 as Histoire naturelle et mythologique de l’ibis which included some very nice illustrations.

from Savigny (1805) hand-coloured by Louis Bouqet

Georges Cuvier, one of the leading French biologists of the day, was asked by Napoleon to join the Egyptian contingent, but he suggested that Savigny go instead, so he could continue his work on molluscs. But it was Cuvier who first measured two mummified birds brought back from Egypt by Col. Jacques François-Louis Grobert [6] from the catacombs at Saqqara.  Cuvier initially concluded that those birds were probably curlews as they were smaller than some contemporary ibis specimens [7]. He later measured two mummies that Geoffrey Saint-Hilaire had brought home from Thebes. Those measurements plus the colours of some intact feathers convinced him that the mummies collected by Saint-Hilaire were indeed sacred ibises, and his 1804 paper has a very nice summary of his reasoning and all of the previous mis-identifications. Cuvier’s assistant even pieced together the bones from different mummies to make a complete skeleton (see picture below).

Even though the ibis mummies from Saint-Hilaire were not exactly the same size as contemporary birds, Cuvier also used those measurements to bolster his arguments of the fixity of species—evidence that species were created once by a deity and did not change through time. This argument put him at odds with his colleague Lamarck who argued that species changed through geological time.

I was made aware of this ibis story in a new essay in PLoS Biology [8], by Caitlin Curtis, Craig Millar and David Lambert. As Jerry Coyne noted in an essay on his Why Evolution is True site, not many evolutionary biologists seem to be aware of this as an early test of evolutionary change. The reason, I think, is that it was not actually a test [9]. The story is actually rather well known and has been published many times in scientific journals and the popular press ever since Cuvier’s initial publications [10]. While the new essay summarizes many aspects of this story the authors present no evidence in support of some of their claims and I am not entirely convinced by some of their assertions.

Cuvier (1804) identifies the mummies as sacred ibises

When interpreting the past here is always a danger of applying present knowledge and values incorrectly. In this case, I cannot yet tell if my different interpretation of this interesting story is correct. I will need to read the work of Cuvier, Lamarck, Saint-Hilaire and Savigny in the original French and Latin to put the whole story in context but that will take a while, even though all of the relevant texts are now available online. I will revisit the topic when I have done the necessary research.

Whether the details in this new essay by Curtis and colleagues are correctly interpreted or not, it does end with a curious conclusion that I feel deserves some further discussion: Of great importance is the reminder, even today, of the power of a strong personality and that the belief in “what they know to be true” can dramatically influence the direction of science and public opinion. I do not think that anyone would dispute that strong personalities and beliefs can influence science and public opinion. Take, for example, Julian Huxley’s rejection of Darwin’s ideas on sexual selection [11], undoubtedly reducing interest in that topic for the next 50 years or so.  And while it is true that Huxley and Cuvier had strong personalities, and were great communicators and relatively powerful men, I think that their arguments held sway largely because they made them clearly and because there was neither compelling evidence nor any clear and logical mechanisms to explain the existing patterns. In both cases the delays in the progress of science were reasonably short and probably needed the ideas and considerable evidence presented by Darwin and Wallace, and Williams and Trivers, respectively, before there could be any real progress.

Finally, it has probably not escaped your notice that the African Sacred Ibis has been depicted on the cover of The Ibis in one form or another ever since 1859. This may seem a bit odd as that bird does not occur in the wild in Britain and only sparsely in southern Europe through introductions. Thus the sacred ibis does not really appear to be a fitting symbol for the British Ornithologists’ Union. There is a long and interesting story there, but that too will have to wait for another day.


  • Birkhead TR, Wimpenny J, Montgomerie R (2014) Ten Thousand Birds: Ornithology since Darwin. Princeton, NJ: Princeton University Press.
  • Cuvier G (1804) Mémoire sur l’ibis des anciens Égyptiens. Annales du Muséum d’histoire naturelle 4:116-135. [available here]
  • Cuvier G (1812) Recherches sur les Ossemens Fossiles de Quadrupèdes : où l’on rétablit les caractères de plusieurs espèces d’animaux que les révolutions du globe paroissent avoir détruites, t1-4. [Studies of the Fossil Bones of Quadrupeds, volumes 1-4] Paris: Deterville. [available here]
  • Cuvier G (1826) Discours sur les révolutions de la surface du globe, et sur les changemens qu’elles ont produits dans le régne animal. Paris: G. Dufour. [available here and in English translation of the 1825 edition here]
  • Lacépède B-G-E, Cuvier G, Lamarck J-B (1802) Rapport des professeurs du Muséum sur les collections d’histoire naturelle rapportées d’Égypte, par E. Geoffrey. Annales du Museum d’Histoire Naturelle 1: 234–241. [available here]
  • Latham J (1790). Index Ornithologicus, Sive Systema Ornithologiae: Complectens Avium Divisionem In Classes, Ordines, Genera, Species, Ipsarumque Varietates (2 Volumes, in Latin). London: Leigh & Sotheby. [available here]
  • Le-Suer RB, ed (2012) Between Heaven and Earth: Birds in Ancient Egypt. Chicago: The Oriental Institute of the University of Chicago
  • Linnæus  C (1758) Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, 10th edition.  Stockholm: Laurentius Salvius.
  • Rosser WH (1837) Mummy of Egyptian Ibis. The Gentleman’s Magazine 6 (new series): 145-148. [available here]
  • Savigny J-C (1805) Histoire naturelle et mythologique de l’ibis [Natural and Mythological History of the Ibis]. Paris: Allais. [available here]


  1. ibis described: see Latham 1790 page 706 where he calls it Tantalus aethyopius
  2. Sacred ibis once common in Egypt: but it is no longer found in that country, disappearing as the swamps and marshes were drained to provide land for the increasing population and agriculture.
  3. Late and Ptolomeic Periods: about 700 BC until 30 BC ending with the death of Cleopatra and the conquest of Egypt by the Romans
  4. ibises eating refuse: in Australia, where they have been introduced, they are often called ‘bin chickens’ as they are often seen foraging in trash cans in city parks. In the park beside the Australian National Museum in Sydney, I once watched a very dirty-looking ibis sneak up behind some picnickers then reach over the shoulder of a little boy to snatch his sandwich out of his hand. Clearly, their bills are adapted for sandwich snatching (!), and they are fearless.
  5. savants: these were scholars and scientists. The Journal des Sçavants (later called Journal des Savants) began publishing in January 1665, a couple of months before the Philosophical Transacations of the Royal Society, considered (erroneously) by many to be the first scientific journal. Frankly, I don’t see that it matters who was first, or even if it was one of those two.
  6. Grobert: (1757-181?) was a French artillery officer who wrote about the pyramids etc. on his return from the Egyptian campaign. See Cuvier (1826), which updates some of the information in his 1804 publication about his later study of some different mummies.
  7. smaller than contemporary ibises: On examining four more ibis mummies, Cuvier recognized that one of them was a juvenile based on its bone structure (Cuvier 1826). As a result, he realized that they may not be curlews at all but simply smaller, juvenile ibises. This is not so surprising as it turns out the many of the ibis mummies were clearly made from juvenile birds. No doubt the priest-farmers who raised the ibises for the votive market saw no reason to keep the birds any longer than was needed to make them suitable for mummification.  That just made good economic sense to maximize their profits.
  8. article in PLoS Biology: unlike the scientific articles in that journal, this one is labelled ‘Essay’ which they say “are opinionated articles on a topic of interest to scientists, as well as to a broader audience, including the general public”. Opinions are fine but I am surprised at the absence of clear evidence in support of the claims made.
  9. not actually a test: Cuvier simply used his measurements identifying the mummies as sacred ibises to suggest that there had not been much change in their morphology in the past 3000 years. But the ibis was just one of many examples that he referred to. Moreover, as Coyne noted, this was at best a ‘one-way test’ as any lack of change would be consistent with slow evolutionary change. Cuvier even acknowledged that the the measurements were not the same between the mummies and contemporary ibises. I don’t see this as a test of any kind because Cuvier was unlikely to be convinced by any such results: if the mummies were the same as extant ibises, then no change; if they were different then they must be different species.
  10. the story of Cuvier’s ibis measurements: in a quick search on the internet, I found more than 20 articles on Cuvier’s ibis measurements dating back to Rosser (1837)
  11. Huxley and sexual selection: see Birkhead et al. (2014)

NOTE As some of you may have noticed, this essay was briefly posted by accident in draft from a week ago. I immediately deleted that version from this blog and the final version above is substantially different, correcting several errors in the original and providing additional information, references and links.

The Invisible Women

BY: Bob Montgomerie, Queen’s University | 3 September 2018

At next year’s annual AOS conference in Anchorage, Alaska, the role of women in ornithology will be one of the highlighted themes. This is an important initiative for several reasons, and will be the focus of several posts here in the coming months.

Most ornithologists are familiar with the names and accomplishments of Margaret Morse Nice, Rachel Carson, Brina Kessel, Fran Hamerstrom, Louise de Kiriline Lawrence, and Florence Merriam Bailey, but what about Hilda Cinat-Thompson [1], Lady Elizabeth Symonds Gwillim, Genevieve Estelle Jones, and Althea Sherman? Women play such a visible role in ornithology (and most sciences) today, that it is easy to forget that women ornithologists were scarce before about 1960. Even those women who contriubuted to the history of ornithology tend to be relatively invisible.

As I have highlighted previously [2], the national ornithological societies that formed in the 1800s were all founded by men, and women were very much in the minority of their membership for much of the twentieth century. That’s just a fact, and I don’t see any point in attempting to rewrite that history. There is a lot to be gained, however, in knowing more about the women who did contribute to the development of ornithology and celebrating their contributions. Unfortunately, the contributions of many of those women to ornithology were never recorded, so they may forever be invisible—at least by name—to history. Today’s post highlights just one of what must be many instances of invisible women who made a great contribution.

Dresser’s Birds of Europe (1871-82) letter bound into 9 Volumes, plus an index and a supplement

From 1871 to 1882, Henry Dresser published the 84 parts of his monumental A History of the Birds of Europe. Dresser was a prominent timber and iron merchant by day, and an ornithologist in the evenings—and presumably weekends and holidays, given his phenomenal productivity. During the mid-1800s, he made his fortune as a merchant in London, and began collecting birds and eggs on various field trips [3]. He eventually amassed a huge collection of bird specimens, eventually purchasing specimens from collectors and dealers around the world. In addition to a handful of excellent books, he also published more than 100 papers about birds. Dresser was well-connected in ornithological circles, regularly corresponding with Alfred Newton and Alfred Russel Wallace, for example, and was BOU secretary from 1882-88 [4].

PartcoverDresser’s Birds of Europe was published in separate parts, by subscription, so that he could use the income from subscriptions to fund the entire project [5]. Subscribers, of which there were eventually more than 300, received an unbound section of both letterpress and plates in blue paper covers every month, and many of those were eventually bound into leather-covered volumes by the subscribers. The whole set cost subscribers £52 10s, or about £5000 (roughly $6500 US in today’s currency).  You can pick up a full leather-bound set today for only $23,000 US at some of the antiquarian booksellers, which is actually quite a bargain given the rate of inflation over the past 140 years [6].

Each of the 634 species in Birds of Europe is illustrated on a superb colour print of the bird—often male and female, sometimes a chick or two—produced mainly by the outstanding 19th century illustrator J. G. Keulemans, plus a few by Joseph Wolf and Edward Neale. These illustrations are remarkable for their accuracy and the pace at which Keulemans made them, often in the midst of working on other projects.

Ruffs displaying from Dresser (1871-82)

To save time (and costs) Keulemans made most of these illustrations by drawing with sharp-pointed greasy crayon directly on the lithographic stone that would be used to make black outline drawings that would be coloured by hand to make the plates. Keulemans was renowned for his ability to use a study skin to make a life-like painting of a bird that he had not even seen in the wild. The fact that he could draw in crayon (in reverse!) on a lithographic stone without working from a sketch seems impossible to me, but then again I have no artistic talents whatsoever. Once the first satisfactory print was made, he used watercolours to make the final master copy. Keulemans only painted the master copy—all of the others that eventually ended up as plates in the book were painted in watercolours mainly by young women [7] in the employ of colourist workshops, using Keulemans’ originals as a guide. We do not know who these women were but the quality and quantity of their work—and thus their contribution to what many consider to be one of the finest bird books ever produced—was outstanding.

telephone switchboard ca 1900

As was the custom of the day, Dresser thanked the men who owned the companies who did the colour work and made no mention of the women who actually did the colouring. I am reminded of an old illustration of women doing all of the work at a switchboard in Paris with the male supervisor overseeing [8]. Here is Dresser, in his Preface: “…and the colouring was entrusted to Mr. Smith and Mr. Hart, the latter of whom is well known as the artist employed by Mr. Gould during the publication of all his later works.” [9].

Lest you think the colourists had no particular talent and were merely making passable copies of the works by the master (Keulemans) have a look at some of the detail, below, on a couple of the plates. Exquisite. The pace at which those women worked must also have been phenomenal. We do not know how many women were employed by Smith and Hart to do the colouring, but we know that they produced a quarter of a million copies (yes, 250,000! [10]) of the Keulemans’ originals in about 12 years. Even at one copy a day—a pace that I cannot even imagine—about 65 artists would have been needed to do all of that colouring [11].


Details of 6 plates in Dresser (1871-96) showing the lovely brush work and attention to detail

Keulemans apparently inspected all of the colouring to ensure accuracy and consistency.  I have looked at several copies of the original plates and cannot detect any difference between copies of the same plate even though they must have been painted by different, unknown, women.

We often vilify the practices of the past because they do not match our contemporary standards of fairness, equality, and recognition. No doubt our own academic descendants will similarly criticize us for our apparent failings. Instead, I think there is some value in trying to identify work that made important contributions to the history of ornithology, even in cases like this where we cannot positively identify who actually did that work. It would be interesting to know if any of the colourists for Dresser’s work went on to be ornithologists or artists in their own right.


  • Dresser HE (1871-82) A History of the Birds of Europe, including all the species inhabiting the Western Palaearctic Region [84 parts; first 13 parts coauthored with RB Sharpe]. London: Privately published. (available online here)
  • McGhie HA (2017) Henry Dresser and Victorian ornithology: Birds, books and business. Manchester, UK: Manchester University Press.


  1. Hilda Cinat-Thompson: did pioneering work on sexual selection in budgerigars but is so little known about her that the only references I can find to her online are in the book I wrote with Tim Birkhead and Jo Wimpenny on the history of ornithology (see here)
  2. formation of ornithological societies: see previous posts here, here, and here
  3. various field trips: to Texas, Mexico and New Brunswick (Canada) for example. Most of his collection eventually went to the Manchester Museum
  4. Henry Dresser’s life: details here were taken mainly from a new book (McGhie 2017) about Dresser that I will be reviewing here in a few weeks
  5. by subscription: the initial subscription price was £6 6s (about $8.50 US) per year for 12 parts with each part containing 10-12 species, and the whole project planned to take 6 years comprising about 72 parts, with each year constituting a volume (McGhie 2017, page 137)
  6. rate of inflation: an online calculator here, suggests that $6500 in 1880 would today be worth $153,000.
  7. we do not know who the colourists were: it might be possible to examine the records from Hart and Smith, and their workshops, to actually identify the colourists but that information is not yet readily available
  8. telephone switchboard operators: many of the earliest switchboard operators were young men, but it was soon recognized that women were generally more courteous. Probably more significantly, though, women were paid at only one quarter of the salary of the men! More info here.
  9. Dresser quotation: from page iv of Vol 1 in Dresser (1871-1882). He is referring here to Smith, Elder and Co., and to William Hart who was both an artist and a colourist who, presumably, supervised the work of several others.
  10. 250,000 copies: actually at least 214,587 coloured plates based on 633 plates per volume and 339 copies at one copy per subscriber. Presumably there were more plates completed than there were subscribers, as the number of subscribers grew through the 12 years of the project.
  11. 65 colourists: based on 250,000 coloured plates, about 250 working days per year, and 12 years for the project

IMAGES: all those of and from Dresser’s Birds of Europe were taken by the author in August 2018 at the Blacker-Wood Collections in the McGill University Library, with thanks to the librarian, Lauren Williams, for permission to use those photographs here; women telephone operators from Wikimedia Commons

Joe Grinnell’s Notes

BY: Bob Montgomerie, Queen’s University | 13 August 2018 (posted 21 Aug 2018)

For at least 400 years, ornithologists—and presumably naturalists of every stripe—have kept notebooks recording each day’s observations from the field. In 17th century England, these were called ‘Commonplace Books’, rather large bound volumes that were used by scholars to record ideas, notes about what they read, experiences and observations. This was the Renaissance, and the beginning of the scientific revolution, where scholars were questioning everything, and basing conclusions on direct observations rather than hearsay, ancient texts, and idle speculation.

Detail from a page in Linnaeus’s commonplace book

John Ray and Francis Willughby [1] each had their own Commonplace Book, as required by their tutors at Cambridge.  In the late 1600s, the great English philosopher John Locke considered Commonplace Books to be so important to the progress of science that he published a scheme for properly indexing a commonplace book in an addendum to his influential An Essay Concerning Human Understanding [2]. And in the 18th century, Linnaeus used his Commonplace Book to record and develop his ideas about his binomial system of nomenclature, resulting in his Systema Naturae [3].

Commonplace books seemed to be de rigueur for scientists and scholars through the 1800s eventually evolving into the specialized (rather than all-encompassing) small notebooks (e.g. Moleskins) and field notebooks (e.g. Rite in the Rain) used by writers and naturalists, respectively, throughout the 20th century.

A page from Grinnell’s Field Journal from the Mojave 1914

In the early 1900s, the American ornithologist Joseph Grinnell thought that field notebooks were so important that he developed a systematic method of note-keeping that he taught all of his students and colleagues. His method, sometimes called the Grinnell System, involves at least two different books—the Field Notebook, carried everywhere to record observations immediately, and the Field Journal, to daily record experiences and observations as in a diary, using the Field Notebook. Each diary-like entry in the Field Journal is written in the evening, using the Field Notebook for details. The Field Journals, or separate notebooks, also include Species Accounts compiled during the course of a field trip, and a Catalog, recording the details of all specimens collected. The method seems simple enough but requires some discipline to maintain during busy field work. Grinnell even went so far as to recommend the sort of paper and ink needed to make the method historically valuable: The India ink and paper of permanent quality will mean that our notes will be accessible 200 years from now….we are in the newest part of the new world where the population will be immense in fifty years at most. [4]


I am an academic descendant of Grinnell [5] and while I am not a very disciplined diarist, I treasure the 57 notebooks that I have used to chronicle my field activities over the years. These books contain some data but they are mostly a summary of where I went, what I did, what the weather was like, who my companions were, what I found interesting each day in the field, and ideas for further work. My field data sheets and recordings occupy another 5 metres of book shelf and a few terabytes of hard drive space.

In 1908, Grinnell was appointed as the first director of the Museum of Vertebrate Zoology at Berkeley, where he set out to build a collection of birds and mammals from California. To do that he embarked on a series of expeditions to the Colorado Desert (1908), The Colorado River (1910), Mount Whitney (1911), the San Jacinto Mountains (1913), the Sierra Nevada (191–1920), and Lassen Peak (1924-1929).

Grinnell kept such careful field notes that the MVZ scientists decided to survey some of those same areas beginning in 2002, to see what, if anything, had changed over the past century. They called this the Grinnell Resurvey Project. Grinnell did not actually conduct censuses using repeatable, modern-day methods, but he did provide enough information that reasonable comparisons could be made.

Earlier this month, PhD student Kelly Iknayan and AOS Past President Steve Beissinger published a paper in PNAS using both Grinnell’s surveys and the recently completed replication to analyze the changes in bird fauna in the Mojave Desert of California. The nice thing about this resurvey is that most of the sites visited by Grinnell in the Mojave are on federal lands, with little or no anthropogenic influence in the intervening 100 years. The results are clear…and depressing.

Surveying 61 of the same sites studied by Grinnell, they found that the number of bird species at each site had declined, often significantly (red dots of figure below). And the number of sites where they found different birds had also declined for >125 of those 135 species. Only the Raven was found at significantly more sites a century later (blue dot, below). IknayanFIGmod

By evaluating several potential causes for these changes, Iknayan and Beissinger found that climate change was the strongest predictor, particularly with respect to increasing drought conditions. As they point out, in their paper’s abstract: Climate change has caused deserts, already defined by climatic extremes, to warm and dry more rapidly than other ecoregions in the contiguous United States over the last 50 years. Desert birds persist near the edge of their physiological limits, and climate change could cause lethal dehydration and hyperthermia, leading to decline or extirpation of some species. [6]

I expect that Iknayan and Beissinger take better field notes that I do, especially as they are both also academic descendants of Grinnell [5] and work in his shadow at Berkeley. But even the best field biologists’ note-taking abilities are rapidly becoming anachronisms, I fear, with the advent of eBird, automated recording devices, and digital database apps. I think this is sad, not because I long for the good old days—I am a quite tech savvy—but because those detailed field journals are an important historical record [7[ that show both the human side of field work and the nuances associated with collecting data.

You could argue that Grinnell’s field surveys would have been more useful today if he had digitized his records and taken more quantitative measures, and you would be right to some extent. But field naturalists a century from now will no doubt lament the passing of the commonplace book and the Grinellian field notebook when they try to understand our quantitive, digitized, data stored faithfully in online repositories if those data are not also supplemented by a little personalized narrative.


  • Charmantier I (2011) Carl Linnaeus and the visual representation of nature. HIST STUD NAT SCI 41:365–404.

  • Iknayan KJ, Beissinger SR (2018) Collapse of a desert bird community over the past century driven by climate change. Proceedings of the National Academy of Sciences 201805123.

  • Locke J (1689) An Essay Concerning Human Understanding. London: The Buffet.

  • Linné CV (1766) Caroli a Linné. Systema naturae : per regna tria natura, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis / (t.1, pt. 1 (Regnum animale) (1766)). Holmiae :Impensis direct. Laurentii Salvii.


  1. Ray and Willughby: see previous posts here, here, here, and here
  2. An Essay Concerning Human Understanding: see Locke (1690) available here
  3. Sytema Naturae: see von Linné (1766)
  4. Grinnell on paper and ink: cited from Wikipedia article on Grinnell, here
  5. Academic descendants: me through Peter Grant to Ian McTaggart-Cowan to Grinnell; Beissinger through Bobbi Low to Frank Blair to Lee Dice to Grinnell (see here and here for details)
  6. Quotation about climate change: from Iknayan and Beissinger 2018, abstract
  7. Important historical record: see here for example

IMAGES: Linnaeus’s notebook from Charmantier (2011); Grinnell’s notebook from the Grinnell ECOREADER;  graphs modified from Iknayan and Beissinger (2018)

A great store of fowle

BY: Bob Montgomerie, Queen’s University | 6 August 2018

Four hundred and eight years ago this month—in August 1610—Henry Hudson and his crew of 21 on the tiny ship DISCOVERY entered Hudson’s namesake bay in search of a northwest passage to the orient. As far as we know, Hudson’s 1610-1611 expedition was the first time that Europeans had recorded the sighting of an identifiable arctic bird on its breeding grounds in North America. Martin Frobisher, for example, had previously visited Baffin Island three times in a vain attempt to mine for gold [1] but he made virtually no note of the birds [2].

A map made in 1612 from Hudson’s surveys on his final expedition

Hudson’s crew famously mutineed in June 1611 after a dreadful winter spent on their ship trapped in the ice of James Bay. The 12 mutineers set Hudson, his son, and 7 loyal  seamen adrift in rowboat and their fate is still unknown [3]. What we do know about Hudson’s final expedition comes from the writings of one of the mutineers, Abacuk Prickett, who wrote about it after returning to England [4]. Prickett was one of the four mutineers who was tried (and acquitted) for the mutiny, and there has always been some suspicion that his narrative was biased in a way that was designed to save him from the gallows. Nonetheless, there is no reason to expect that his account of the birds is not as accurate as could be expected for a document being written, we presume, largely from memory.

Prickett records that their first landfall in the Canadian arctic was in July 1610 on the ‘Iles of Gods Mercie’, probably the islands off the south coast of Baffin Island [5] near the present-day settlement of Kimmirut (formerly Lake Harbour) in Nunavut. There, they “sprung a covey of partridges which were young: at the which Thomas Woodhouse shot, but killed only the old one” [6]. Given the current breeding ranges of the two arctic ptarmigans, these were almost certainly Rock Ptarmigan, which makes it the first bird species recorded in Arctic North America and, fittingly, the official bird of Nunavut.

Their next landfall was at Digges Island [7] on 3 August. A small crew went ashore, including Prickett who said “In this place a great store of fowle breed…” [8], almost certainly referring to the huge colony of Thick-billed Murres nesting on the cliffs there, today numbering some 300,000 breeding pairs.

Clets on St Kilda

On Digges, Prickett also noted that “Passing along wee saw some round hills of stone, like to grasse cockes, which at the first I tooke to be the worke of some Christian. Wee passed by them, till we came to the south side of the hill we went unto them and there found more; and being nigh them I turned off the uppermost stone, and found them hollow within and full of fowles hanged by their neckes.” [8]. What he is referring to here are small domed stone huts, about 2 m in diameter, built by the local Inuit to hang, dry and protect their game from predators.

Remarkably, my colleague Tony Gaston, who studied the murres on Digges in the 1980s, found at least four of the same drying huts described by Prickett. As Gaston noted, these are very similar to a structure called a ‘clett’ (also ‘clet’) that the inhabitants of the Outer Hebrides use to dry and cure fish and birds (see photo).

From Digges, the explorers headed south, ecstatic that they might have found the passage to China, as winter approached. By the time they reached James Bay, they knew that there nowhere near the orient. But on  10 November DISCOVERY was trapped in the sea ice so the crew prepared for the winter. During that harsh winter, they often went ashore to hunt, taking as many as 1200 ptarmigan, enough for each man to have one to eat every day or two for three months: “for the space of three moneths wee had such store of fowle of one kinde (which were partridges as white as milke) that wee killed above an hundred dozen, besides others of sundry sorts…The spring coming this fowle left us, yet they were with us all the extreame cold. Then in their places came divers sort of other fowle, as swanne, geese, duck, and teale, but hard to come by.” [9]

Digges Island 1952 photo by Les Tuck

With the ship freed from the ice, the mutineers set Hudson and the others adrift at the top of James Bay in June 1611, and headed back to Digges to stock up on murres and their eggs for the trip home. There, they encountered a band of the local Inuit collecting eggs and catching adult murres with a noose, much the same way that today’s researchers catch murres for banding: “Our boat went to the place where the fowle bred, and were desirous to know how the savages killed their fowle: he shewed them the manner how, which was thus: they take a long pole with a snare at the end, which they put about the fowles necke, and so plucke them downe. When our men knew that they had a better way of their owne, they shewed him the use of our peeces, which at one shot would kill seven or eight.” [10]

The natives became frightened and suspicious of the mutineers, attacking an unarmed party that had gone ashore one day to shoot some murres. Three of that party were killed but the others escaped. The remaining mutineers went to another part of the colony where they shot enough birds to (barely) get them home.

None of these vague observations of birds by Prickett really made any useful contribution to ornithology, and I tell this story mainly as an introduction to the history of ornithology in the North American Arctic. By the late 18th century, explorers and naturalists were making regular forays into what is now Nunavut, the Northwest Territories, Yukon, and Alaska. Those later expeditions did make many useful contributions to ornithology, finding the breeding grounds and documenting the breeding biology of many Arctic birds for the first time.

coversSome of this early Arctic ornithology is described in a forthcoming 2-volume book on the Birds of Nunavut that will be launched at the upcoming IOC meeting in Vancouver. I wrote the history chapter for that book, but the limitations of space meant that many stories, images, and details had to be left out. As for much of the history of ornithology, this blog provides a unique opportunity to expand on the details of scholarly books and papers, as I have done here with the story of Abacuk Prickett.


    • Collinson R, editor (1867). The Three Voyages of Martin Frobisher, in Search of a Passage to Cathaia and India by the North-West, A.D. 1576-8. London: Hakluyt Society. [available here]
    • Gaston AJ, Cairns DK, Elliot RD, Noble DG (1985) A natural history of Digges Sound. Canadian Wildlife Service Report Series 46:1–63.
    • Mancall PR (2009) Fatal Journey: The Final Expedition of Henry Hudson. New York: Basic Books.
    • Prickett A (1860). A larger discourse of the same voyage, and the successe thereof. In G. M. Asher (Ed.), Henry Hudson the Navigator: the original documents in which his career is recorded (pp. 98-36). London: Hakluyt Society. [available here]
    • Richards JM, Gaston AJ, editors (2018) Birds of Nunavut. Vancouver: UBC Press.


    1. Frobisher mining for gold: on his third expedition in 1578, for example, he took back to England 1350 tonnes of ore from the vicinity of Iqaluit (formerly Frobisher Bay) only to discover when he got back to England that the ‘gold’ was iron pyrite. No doubt he felt like a fool.
    2. Frobisher’s birds: Collinson (1867) has only three mentions of birds (fowle) in Frobisher’s writings and these were all with reference to birds and eggs being caught by the natives for food. It is impossible to know what birds he was talking about.
    3. Hudson’s fate unknown: there is speculation, however, that the men made their way south where were taken captive by the natives, then transported to the vicinity of Ottawa, Ontario (see here for details)
    4. Prickett’s account of the expedition: see Prickett (1860), in a volume by the Hakluyt Society, established in 1846 to publish original accounts of voyages of discovery. Prickett’s account was actually first published in 1825. Prickett is often spelled ‘Pricket’ but I am using the spelling on his account in the 1860 volume.
    5. Iles of God’s Mercie: these are shown on Hudson’s map (above), offshore where he labels ‘Goods Merces’
    6. Quotation about partridges: from Prickett 1860 page 103
    7. Digges Island: Hudson named this Deepes Cape, thinking initially that it was part of the mainland
    8. Quotations about ‘fowles’: from Prickett 1860, page 107
    9. Quotation about hunting birds in winter and spring: from Prickett 1860, page 113
    10. Quotation about Inuit method of catching murres: from Prickett 1860, page 128

IMAGES: Hudson map from the frontispiece of Asher (1860) where Prickett’s account was published; Clets on St Kilda from Wikimedia Commons; Digges Island photo by Leslie M. Tuck in the author’s collection; book cover from UBC Press.

But is it art?

BY: Bob Montgomerie, Queen’s University | 23 July 2018

LSNYBig cities—maybe surprisingly—are good places to live if you are interested in natural history. While country folk have nature on their doorstep, city folk have to work a little harder to find woodland and wildlife. The advantage to city life, though, is that there are lots of like-minded people nearby and the resources for nature study and discussion can be amazing—museums, aquaria, naturalist clubs, universities. The Linnaean Society of New York, for example, has brought together naturalists from all walks of life since 1878 to meet regularly at the American Museum of Natural History and to hear lectures from world-class naturalists and biologists. And just across the road from the museum lies Central Park, not a bad place for birding.

When I was 11 years old, my parents sent me off each Saturday morning to the Royal Ontario Museum (ROM), a few blocks from our home in downtown Toronto, to meet with 250 other children and young teens in the Toronto Juniour Field Naturalists’ Club TJFN). This was a magical experience for me, getting to hang out for 3 hours every week with like-minded people, hearing talks and watching natural history films, getting to explore behind the scenes in a great museum, and regularly going on field trips to nearby parks and conservation areas inside or very near the burgeoning city.

The TJFN was organized into about dozen special interest groups—mammals, birds, insects, and the like—that we broke into after the initial presentation to share our experiences and learn from the group leader. I first joined the mammal group, led by a young man named  Bob, a local school teacher who had just returned from a 14-month-long expedition from Africa to Australia by Land Rover with one of his friends. That expedition was what our dreams were made of, and my three naturalist buddies and I spent many an evening planning to make a similar trek one day by Land Rover from Pont Barrow to Tierra del Fuego. Bob taught art at a local high school and gave us budding wildlife artists some great feedback on our feeble attempts to draw the local animals and plants. Years later he developed into one of the world’s best and best-known wildlife artists—Robert Bateman.

Ruby-crowned Kinglet by Bob Bateman, age 17

Bob was certainly the first real artist that I had ever met, but those TJFN meetings eventually brought me into contact with the ROM’s excellent artists and illustrators—Terry Shortt (birds), Paul Geraghty [1] (mammals, birds and dinosaurs), Peter Buerschaper (fishes) and Anker Odum (insects). It was only years later that I came to appreciate what a privilege it had been to know such outstanding wildlife artists.

In the late 1960s when I was working at the ROM for a year, I also got to meet a slew of young wildlife artists who brought their work into the bird department both to get some feedback on their latest work and to borrow skins to help them with details of plumage and bare parts. Names like J. Fenwick Lansdowne, Glen Loates, Barry Kent Mackay, and George MacLean might be familiar to some readers of this blog.

Once, on a long drive from Toronto to Thunder Bay, Paul Geraghty and I had a lively discussion about the nature of bird ‘art’ and bird ‘illustration’. I naively considered any drawing or painting of a bird to be ‘art’ but Paul, quite rightly I think, made a distinction between the two. To him, birds drawn or painted to highlight the bird’s features for the purpose of identification or to enhance some text were illustration, and birds drawn or painted in such a way that the work evoked a more emotional response were art.

For years after that trip, we often debated about paintings of wildlife. When I told Paul about some new painter I had discovered, he would often say, “but is it art?” I think we agreed that there was no point to trying to make a clear distinction along what was clearly a continuum, but the ends of that spectrum were quite clear to us—Bruno Liljefors’s painting of a Golden Eagle was art, but Roger Tory Peterson’s painting of that species in his field guide was illustration.

Golden eagle by Bruno Liljefors (L) and Roger Tory Peterson (R, bottom)

I think it’s fair to say that Audubon was among the first [2] to blur the distinction between art and illustration. Before Audubon, birds were (usually) painted [3] as part of a scene, portrait, or still life (art), or in bird books to show the salient features of a species (illustration). Most of Audubon’s portraits in his Birds of America served both purposes, often being very ‘artistic’ in composition, depicting birds in habitats, often flying, preying, defending or courting. My favourite Audubon (below) certainly blurs the lines for me, combining flight, attempted predation, and accuracy useful enough for identification.

Bobwhite and red-shouldered Hawk watercolour by JJ Audubon ca 1825

Before Audubon, most bird illustration looked wooden [4], showed little or no action or habitat, and lacked the sort of ‘jizz’ that makes a painted bird look real. Bird art, of course, goes back far into prehistory with ancient depictions of loon-like birds found in archaeological digs, and myriad birds in Japanese, Chinese and other works prepared before the start of the Christian Era. Some of these early bird drawings and paintings may have been used for ‘identification’ but my guess is that they were mostly symbolic or ornamental.

Unequivocal bird illustration may have begun with Frederick II in his manuscript De Arte Venandi cum Avibus where birds were drawn in colour in the margins to enhance the accompanying text. Frederick did not identify his birds explicitly but, in his 1983 paper, William Yapp made a good stab at figuring our which species were being illustrated. The bird drawings in Frederick’s manuscript represent at least 60 species and are remarkably identifiable today. OK, I said ‘unequivocal’, but you could argue that they are really art because of their composition, and the depiction of habitats and behaviours, much as Audubon did 600 years later.

bottom of folio 7r in Frederick II’s De Arte Venandi cum Avibus (ca 1245)

Willughby and Ray’s Ornithology may have been the first serious attempt to accurately depict birds in a book [5], showing their ‘characteristic marks’ for identification. Ray, however, was not pleased with result and the illustrations do look very wooden by today’s standards. For the next 150 years or so bird illustrations in books became progressively more life-like in part due to improvement in printing methods—certainly birds in ‘art’ were remarkably accurate more than 2000 years ago.

Possibly due to Audubon’s influence and success, bird illustrations in books became much more ‘artistic’ in the 19th century with people like George Lodge, Archibald Thorburn, John Gould and many others putting more life, action and habitat into their work for bird books. While there were dozens of great bird artists in the century after Audubon, it seems to me that wildlife art became a ‘thing’ only in the 1970s, possibly fuelled by the critical and commercial success of Robert Bateman.

Snow Geese by Paul Geraghty ca 1973

We ornithologists are very lucky to have attracted such a wealth of talent to our discipline, adorning our books (and homes) with great images of our favourite animals. When Tim Birkhead, Jo Wimpenny and I were putting together our Ten Thousand Birds book on the recent history of ornithology, we decided to start each chapter with an example of superb bird art or illustration produced since Darwin. We were able to include the work of 12 great artisans but we were both delighted and perplexed by the wealth of available material. In the end, we had to make some rather arbitrary choices of whose work to include.

As one example of the current wealth of bird art, you need only attend the annual Birds in Art exhibiton at the Leigh Yawkey Woodson Art Museum, in Wasau, Wisconsin. This year, that exhibition will run from 8 September until 25 November and feature the work of more than 60 outstanding bird artists, including a few names that I recognize, like Tony Angell, Robert Bateman, Guy Coheleach, and Maynard Reece. There are dozens of such exhibitions every year in both North America and Europe, attesting to the popularity of this wildlife art. Art or illustration?…whatever you want to call it, the work is fabulous.


  • Audubon JJ (1827-1838) The Birds of America. Edinburgh & London: J. J. Audubon.

  • Birkhead TR, Wimpenny J, Montgomerie R (2014) Ten Thousand Birds: Ornithology since Darwin. Princeton, NJ: Princeton University Press.
  • Frederick II (~1245) De arte venandi cum avibus. Vatican City, Biblioteca Apostolica Vaticana, codex Ms. Pal. Lat. 1071.
  • Johnsgard PA (1974) Song of the North Wind: a story of the snow goose. U of Nebraska Press.

  • Ray J (1678) The Ornithology of Francis Willughby. London: John Martyn.

  • Yapp WB (1983) The illustrations of birds in the Vatican manuscript of De arte venandi cum avibus of Frederick II. Annals of science 40:597–634.


  1. Paul Geraghty: not to be confused with the British author and illustrator of children’s books, often about dinosaurs and other animals. The Paul Geraghty I am referring to now lives in Saskatchewan and some examples of his early work can be seen in Johnsgard (1974). I have reproduced one of those drawings at the bottom of this essay. The cover of Johnsgard’s book says ‘Illustrations by Paul Geraghty’  but even by Paul’s own restrictive definition I would certainly call this ‘art’.
  2. Audubon among the first: I am being especially cautious here, and throughout this essay, because (i) I am no expert, and (ii) people are passionate about their favourite bird artists/illustrators and there is bound to be lots of disagreement. I do not see much point in playing the ‘who’s the first/who’s the best’ game, but I do think that the diversity of opinions is both interesting and productive. Feel free to weigh in on this topic in the comments.
  3. painted and drawn: I do recognize that these are not the only visual art forms but those are the ones I am dealing with in this essay. I will discuss sculptures, movies, and music some other time.
  4. bird illustrations looked wooden: it is pretty clear that the quality of bird illustration lagged behind that of plants, for example, because birds could not be accurately drawn from life. Bfore the ready availability of binoculars, telescopes and phoitography, birds could not usually be approached and opbserved very closely. As a result, most illustrators worked from dead birds, including Audubon. Audubon, however, often mounted his bird carcasses using wires etc into more or less lifelike poses as models for his work.
  5. Willughy and Ray illustrations: several previous authors (Aldrovandi, Belon, etc) had included bird illustrations in their books but Ray and Willughby made an explicit attempt to be accurate and to include some features useful in identification.

Being Francis Willughby

BY: Bob Montgomerie, Queen’s University | 16 July 2018

Last week I posted a list of recently published books relevant to the history of ornithology. This week I will begin to review them, one at a time, as I finish reading each one. I am a slow reader, so don’t expect me to finish before the end of the year.

This first review is about a very recent book by my friend, colleague and collaborator, Tim Birkhead, at the University of Sheffield. Tim and I have been friends for 30 years, so this review comes to you with a host of disclaimers and biases [1]. One distinct advantage of writing a book review on a blog—rather than for a scientific journal or the popular press—is that I can write in a less formal, more intimate, and ultimately, I hope, more interesting format.

So, here’s a book published in May 2018, by Bloomsbury, who have published some of Tim’s other recent books, as well as the entire Harry Potter series.

WMWBirkhead TR (2018) The Wonderful Mr. Willughby. London: Bloomsbury

About 20 years ago, Tim wrote to me to ask who I thought was the first real ornithologist. He knew that I knew that Aristotle, Frederick II, and Aldrovandi (and others) had written about birds a long time ago, but he wanted to know who I thought had actually written about ornithology in a way that provided the foundation for development of our discipline. 

I guessed John Ray, who had written Ornithologia Tres Libris in 1676 and that was exactly what Tim was thinking, too. But I think he was more than a little surprised that I had even heard about that book.

I had actually spent a couple of hours with the English edition of Ray’s Ornithologia in the rare book collection at McGill University in 1974. I started my PhD at McGill in the fall of 1973 and soon began spending every Friday morning in their fabulous Blacker-Wood Library of Ornithology and Zoology. There weren’t many of us interested in birds at McGill so I usually had the place to myself and got to know the new Blacker-Wood librarian, Eleanor MacLean, quite well. One day she asked if I’d like to see the rare book collection and that’s where I saw 3 copies of the English edition of Ray published in 1678, one of which had all the birds in colour.

Ray1678coverWhen, more than 25 years later, I told Tim about this experience, he thought I must have been mistaken about the coloured edition because the book was not hand coloured. When we visited the library in 2008, sure enough both the hand-coloured book—and Eleanor MacLean—were still there. That unique copy of this book was ostensibly given to Samuel Pepys by Ray himself, and had been purchased by the library’s founder, Casey Wood, in 1922 [2].

Tim was convinced that Ray’s book was an under appreciated masterpiece, so he used it as the foundation for his first book on the history of ornithology *The Wisdom of Birds*, the title of which is a paean to Ray’s best-known work, *The Wisdom of God* [3].

Tim was also convinced that Francis Willugby’s contributions to the book were also under appreciated, to the point that Willughby was virtually unknown, and largely ignored in the history of science in general and ornithology in particular. Willughby and Ray were friends and collaborators but Willughby died when he was only 36, before he had published anything about birds, leaving Ray to write up his research on natural history, comprising books about birds, fishes and insects based on their 20 years of study and exploration focussed on natural history.

Ray published the Latin edition of the book giving full credit to Willughby’s contributions in the preface, but—possibly feeling a bit guilty—put Willughby’s name in the title of the English edition The Ornithology of Francis Willughby. 

In retrospect that title says it all to me—this is Francis Willughby’s contribution to ornithology, written by his collaborator John Ray after Willughby’s untimely death. Many others seemed to have missed that point, Tim told me, giving full credit to the ‘genius’ John Ray and relegating Willughby to the dustbin of history. 

In 1943, the Reverend Charles Raven seemed to have put the final nail in Willughby’s coffin with his detailed and fawning biography of Ray, largely dismissing Willughby as a helper who died before he could make any useful contribution.

To learn more about the relatively unknown Willughby, Tim secured funding from the Leverhulme Trust (UK) to assemble an international coterie of scholars with expertise in the life, times and interests of Francis Willughby. The first product of that collaboration was a scholarly, edited volume on Willughby pulling together all of the known facts [4]. Even though Willughby’s notebooks and archives have largely disappeared in the almost 350 years since he died, there was much to be learned, and that volume is comprehensive. Tim’s new book is based on that project, bringing Willughby to life for a more general audience, and much, much more.

Pepys coloured Orn pl XXWillughby and Ray met at Trinity College, Cambridge, in 1653, where the 18-year-old Willughby was a student and Ray—some 8 years his senior—was his tutor. They seemed to hit it off right away as they shared a passion for natural history and became friends despite the large gap in their social status—Willughby was wealthy and Ray a poor blacksmith’s son. Together they explored the local area with the occasional trip farther afield in southern England, with Ray focussing on botany and Willughby on zoology. But general ideas about natural history, the demarcation and identification of organisms, and thoughts about how to group them logically were their common ground.

With the encouragement and assistance of several friends, Willughby and Ray sought to develop a universal language of words and symbols to describe any organism [5], to compile a description of all of the known species in several large groups [6], and to formulate a classification system that would group together similar species in a hierarchical fashion [7].

On completing their studies at Cambridge, they, and two friends, embarked on a long tour of Europe, starting in the Low Countries. Travelling on foot, horseback, and, presumably, horse-drawn carriage, they gradually worked their way through Germany, Italy and Spain, before returning home. Along the way they explored woodlands, fields and waterways, haunted local markets for exotic species, visited learned naturalists and inspected their collections, and bought drawings and paintings of animals and plants whenever they could. They returned to England [8] with a treasure trove of material and set to work right away to get it organized with a goal to publish on each major grouping.

When Willughby died in 1672, they still had not written anything publishable, but they must have been close to putting pen to paper. Ray immediately began preparing a book on ornithology based on Willughby’s notes and ideas. Willughby, for example, hit on the idea that each species must have distinctive ‘characteristic marks’ that could be used to identify it conclusively, years before Roger Tory Peterson used this principle in his wildly successful series of field guides. I assume that Ray started with birds as a memorial to his friend and benefactor’s too-short life. Within two years of publishing the original book in Latin, he produced an expanded English edition, presumably to reach a wider audience, and, this time, giving proper credit to Willughby in the title. Among his many other works, Ray eventually wrote up Willughby’s notes on the fishes and insects as separate books, both in Latin and both only acknowledging Willughby’s help, as he had in the Latin Ornithologia.

To make the best of the limited information available on Willughby, and to make his new book more interesting and accessible, Tim infuses the text with his own experiences that parallel Willughby’s. Like Willughby, Tim dissected some of the same birds so that he could see first-hand what Willughby must have experienced. Tim has also been to many of the places where Willughby travelled both in England and Europe, read many of the same books, and studied many of the same birds in the field. In a few places he describes some of Willughby’s experiences in an attempt to recreate Willughby’s voice and experience—in essence, being Francis Willughby—to enliven the text. The result—history, biography, ideas and birds—is, in a word, wonderful.


  • Birkhead TR (2008) The wisdom of birds: an illustrated history of ornithology. London: Bloomsbury.
  • Birkhead TR, editor (2016) Virtuoso by Nature: The Scientific Worlds of Francis Willughby FRS (1635-1672). Leiden: Brill
  • Birkhead TR, Charmantier I, Smith PJ, Montgomerie R. (2018) Willughby’s Buzzard: names and misnomers of the European Honey-buzzard (Pernis apivorus). Archives of Natural History 45: 80-91.
  • Montgomerie R, Birkhead TR. 2009. Samuel Pepys’s hand-coloured copy of John Ray’s ‘The Ornithology of Francis, Willughby’ (1678). Journal of Ornithology 150:883-891.


1. disclaimers and biases: as Tim developed this project we discussed it often and I read some bits of the manuscript for him as the writing progressed. We also wrote two papers Germaine to the topic (Montgomerie and Birkhead 2009, Birkhead et al. 2018) but I was not part of the Leverhulme Trust project nor had I seen most of this book before beginning to read it about a month ago.

2. hand-coloured edition: see Montgomerie and Birkhead (2009) for more details

3. Wisdom of God: this is considered by many (non-biologists) to be Ray’s most important work, in which he interprets the natural world as evidence of God’s design. Such an interpreation was the goal of natural philosophy in the 1600s and Ray might well be considered to be the finest example of that approach

4. edited volume on Willughby: see Birkhead (2016) 

5. universal language of natural history: they were encouraged in this endeavour mainly by their friend John Wilkins, but like the much later attempt by others to invent a universal language—Esperanto—this was a complete failure and they soon abandoned the idea, much to our benefit today.

6. descriptions of species: they focussed largely in birds, fishes, insects and flowering plants. There existed previous descriptions (Aldrovandi, Belon, etc) but these tended to be of local fauna, were mainly focused on external features, and could rarely be used for positive identification. Willughby and Ray, on the other hand, recognized that internal anatomy, habitats, song, and behaviour could all be used to distinguish species. 

7. classification systems: while still crude by today’s standards, Linnaeus acknowledged their work in developing his own enduring system a century later. Willughby and Ray, for example, started by dividing all birds into land birds and water birds, then within each group made smaller grouping based on bills, feet, internal anatomy and size.

8. returned to England: Willughby returned to the family estate at Middleton Hall, and provided the funds for He and Ray to devote full time to their natural history work, even after Willughby died.

Summertime and the Birdin’ is easy

BY: Bob Montgomerie, Queen’s University | 9 July 2018

Most of my birder friends don’t do much birding in the summer unless they are involved in breeding bird surveys. Once the flush of spring migration, Global Big Days, and the frenzy of territory establishment have passed, most of them spend the summer months from mid-June to mid-August catching up on their reading, bringing their e-bird lists up-to-date, and planning birding trips for the fall.

Acadian Flycatcher nest & eggs, Rondeau Prov Park, Ontario, 1969

It wasn’t always like this. For many bird enthusiasts, the summer months were the most exciting, because that’s when birds were nesting and egg-collecting was an all-consuming hobby. Even as recently as the 1970s, my old friend George Peck [1] and I spent most of our summer weekends haunting the woods and fields around Toronto in search of nests and eggs to photograph. George was what I might call a reformed öologist—an egg collector—who turned his attention to photographing rather than collecting birds’ eggs when that hobby became not only illegal [2] but scorned and prosecutable in the 1960s. George was a professional veterinarian who was well aware that prosecution for egg-collecting would destroy his career.


When I first met George in the mid 1960s he still had his boyhood egg collection, as it was still legal to possess one then, even though you could not legally collect wild birds’ eggs. With the advent of Kodachrome II and decent colour photography George made it his goal—his life list, if you will—to photograph the nest and eggs of every North American breeding bird, and to building the Ontario Nest Record Scheme into one of the largest and most accurate records of nesting birds ever compiled. George called himself a nidiologist, a term I never hear anymore.

Back in the day—as in the late 1800s—hundreds, no thousands, of men and boys (rarely women) would spend their spare time in summers hunting for birds’s nests and collecting eggs, for fun, for profit, or for science. Some wealthy men—like Walter Rothschild and Johnny Dupont—made huge collections that became the nucleus of many of the large collections in museums today. 

coverAnd there was money to be made because often the wealthiest of collectors did not go into the field at all, but amassed their collections through barter and purchase. For some men, egg collecting was an important source of seasonal income, and thousands of eggs were bought and sold both in personal transactions and by dealers. One dealer, Watkins & Doncaster [3], in 1900, would sell you a Golden Eagle egg for 18/6 ($119.64 in today’s $US), or a Honey Buzzard egg for 7/0 ($45.36 today) [4]. Even the egg of a common British garden bird like the Blackbird would cost 7d (54 cents). As you might expect, price was driven by supply and demand, and demand was driven by the rarity of the bird and the egg pattern [5]. Even given the vendor’s markup, a man could make a decent wage collecting birds’ eggs during the summer.



I would never advocate a return to egg-collecting as a hobby or a vocation, but as I have mentioned before, the great—and scientifically important and useful—egg collections of the world have stagnated, having added precious few specimens for decades. Many of them are also poorly curated, protected, and catalogued, though recently I have seen some  renewed interest on the part of museum curators.

As a working scientist, I can’t even watch birds or record their songs without approval from our Animal Care Committee, let alone find nests and photograph eggs. The general public, of course, is not so restricted, but there is little amateur interest in nests and eggs anymore. Done carefully, and maybe under permit, there would seem to be some value in a renewed interest in nidiology, but that might be too fraught with conservation issues to be very attractive to most people.

There are, of course, always books to read in the summer, and this year there is a superb crop of books for those interested in reading about birds. I have the following pile of books relevant to the history of ornithology on my desk, and will write reviews of most of them in the coming weeks. For now, just a brief description of each book is about.

  • WMWBirkhead TR (2018) The Wonderful Mr Willughby: The first true ornithologist. London: Bloomsbury. [Francis Willughy and John Ray tried to revolutionize natural history in the 17th century. Their classic Ornithologia Tres Libris was really the first encyclopedia of ornithology, with detailed description of all the species known to them. Willughby died when he was only 36, so Ray wrote up all of their findings in classic works on ornithology, fishes and insects. Ray got most of the glory….until now]
  • Brunner B (2017) Birdmania: A remarkable passion for birds. Vancouver: Greystone Books. [A somewhat eclectic compilation of interesting stories about some of the characters that populate the history of ornithology.]
  • Johnson KW (2018) The Feather Thief: Beauty, obsession, and the natural history heist of the century. London: Hutchinson. [The intriguing story of Edwin List who stole valuable bird specimens from the British Museum to get feathers to make expensive flies for fishing]
  • dresserMacGhie HA (2017) Henry Dresser and Victorian Ornithology: Birds, books and business. Manchester: Manchester University Press [While the focus here is on the life of Henry Dresser, from Manchester, UK, this book is a superb window on the state of ornithology in the late 1800s]
  • Olina GP (2018) Pasta for Nightingales: A 17th century handbook of bird-care and Folklore. New Haven, CT: Yale University Press [this is the first English translation, by Kate Clayton, of one of the classics of early ornithology written ins 1622. Replete with contemporary watercolours from Olina’s day.]
  • skelZalasiewicz J, Williams M (2018) Skeletons: The frame of life. Oxford: Oxford University Press [Despite the cover photo, there is not much in this book about birds, but what there is is fascinating, and nicely places birds in the evolution of skeletons. I have already reviewed this book for Times Higher Education in the 14-20 June 2018 issue]


  1. George Peck: was mentioned in my previous posts here, here and here
  2. egg-collecting illegal: In the UK the Protection of Birds Act of 1954 made the colection of birds’ eggs illegal. In tNorth America, that protection began with the Migratory Birds Treaty Act of 1918, but egg collecting continued largely unprosecuted until the UK act of the 1950s. The history of these laws and their enforcement is definitely complex and will be the subject of a later post
  3. Watkins & Doncaster: established in 1874, is still in business, though they no longer sell birds’ eggs. They moved from their location on The Strand in London in 1956, and are now in Hertfordshire (and, of course, on the internet)
  4. egg prices: are listed as shillings/pence in their catalog. I used this site to convert those amounts to today’s currency.
  5. rarity of egg pattern: see here for my previous post on an interesting and rare egg pattern

Much Ado About a Cockatoo (reposted)

BY: Bob Montgomerie, Queen’s University | 2 July 2018

For the past week or so the internet has been abuzz about a cockatoo depicted 4 times in the margins of Frederick II’s De Arte Venandi cum Avibus written around 1245 CE [1]. The story is that this bird suggests a mediaeval trade route from Australia to Italy, overturning the Eurocentric notion that Australia was a dark continent until ‘discovery’ by Dutch sailors early in the 17th century. This story has already been retweeted about 2000 times on Twitter, and has appeared in the popular press worldwide, including CNN, Reuters, ABC (Australia), Japan Times, and The Guardian [2].

Marginal cockatoos (numbers indicate folio pages)

This story began with an original paper published last month  in the journal Parergon [3] by Heather Dalton, Jukka Salo, Pekka Niemelä and Simo Örmä. That paper is wonderfully detailed about the creation and provenance of De Arte, about the source of Frederick’s cockatoo, and the details of the four coloured drawings of the cockatoo in the surviving copy of the original manuscript housed in the Vatican Library in Rome [4]. The popular press has been remarkably accurate in reporting the details of that paper, avoiding the hyperbole and small misleading errors that too often characterize science journalism.

Here, in a nutshell, are 5 quotes that summarize what I think are the important features of this story:

  1. Frederick_II_and_eagle
    Frederick II

    “Frederick II of Sicily made contact with the Kurdish al-Malik Muhammad al-Kamil in 1217… The two rulers communicated regularly over the following twenty years, exchanging letters, books and rare and exotic animals….[like] the Sulphur-crested or Yellow-crested Cockatoo the sultan sent Frederick.”  [5]

  2. “De arte was written in Latin by Frederick or a scribe under his direction between 1241 and 1244…Amongst the nine hundred marginal illustrations of birds, animals, falconers, perches, and falconry equipment are four coloured drawings of the white cockatoo gifted to Frederick II.” [5]
  3. “Discovery of earliest European depiction of cockatoo in medieval book rewrites history of global trade” [6] see also quote 7
  4. “Because the four images in the Vatican manuscript have rarely been reproduced in print, few people are aware of their existence. This may be because many scholars have relied on Casey Albert Wood and Florence Marjorie Fyfe’s 1943 English translation of all six books of the De arte. 11 Although Wood and Fyfe included many illustrations from the Codex Ms. Pal. Lat. 1071, they did not include those of the cockatoo. [5]
  5. “Bearing in mind the shape of the crest, the blue/grey of the periophthalmic ring and the lack of a yellow tinged ear patch, Frederick’s cockatoo was in all likelihood a Triton Cockatoo…or one of the three subspecies of Yellow-crested Cockatoos that have a yellow crest.” [5]
  6. “The main significance about it is we tend to think of our region, not just Australia, but the islands around it, as the very last things to be discovered; the European view is it’s almost this dead continent and nothing was happening until Europeans discovered it.” [7]

While Dalton and colleagues have done a great job summarizing all of the details in their paper, I do have a few quibbles with the final four points listed above. I was, for example, a little surprised to hear of the ‘discovery’ of these cockatoo drawings because I certainly knew about them. As so often happens, I wondered if I had simply failed to realize their significance.

But as the paper so nicely summarizes, previous authors [8] had written about the cockatoos, so this new work might be better characterized as a re-discovery. In fairness, Dalton and colleagues never claimed this to be a new discovery but this is the way that most of the popular press has characterized their work.

Wood and Fyfe p 38: cockatoo is at top of right margin

They do claim (point 4), however, that modern scholars are generally unaware of these illustrations, largely because everyone reads Wood and Fyfe’s translation from 1943. But that’s how I knew about the cockatoo—one of the pages (folio 18v) with the cockatoo in the margin is reproduced, albeit in black and white, on page 38 in Wood and Fyfe’s book. In the caption they even say  “also containing the reference to the parrot (?) sent to Frederick by the Sultan of Babylon”, and in a footnote on page 59 they say it was likely a cockatoo from the Sunda Islands.

Dalton and colleagues do a really nice job of describing the four coloured marginal drawings and they use those details to try to identify the bird. They make the reasonable conclusion that it was likely a female Sulphur- or Yellow-crested Cockatoo. Others have suggested that it might be a White Cockatoo [8]. Nonetheless, I cannot agree with the authors’ assumption that those descriptions are very useful for identification.


For example, in the De Arte illustrations, it looks like the birds have yellowish flanks and back, whereas none of the potential cockatoos have that colouring [9]. They also interpret the shape of the crest as ruling out the White Cockatoo, but the shape and size of the bill, feet, and wing feathers are so inaccurate that I would not be inclined to assume that the crest is correctly drawn. Moreover, none of the illustrations in De Arte show the characteristic yellow cheek patch of the Yellow- and Sulphur-crested.

peacockFiiThe marginal drawings are, after all, crude by modern standards, as you can see from the reproductions above. The manuscript has at least 7 marginal drawings of peacocks [10], for example, that show that, although the artist was remarkably good for his day, he was no Lars Jonsson. Thus it would be entirely reasonable for Frederick’s bird to be a White or Yellow-crested Cockatoo, from Sulawesi (one of the Sunda Islands) or the Moluccas [11]. If that is correct then I am not so sure that this cockatoo really tells us anything about mediaeval trade routes.

It has long been known that there was extensive trade between southeast Asia and the Middle East along the Silk Road, beginning centuries before Frederick’s day. That ‘road’ includes several marine routes extending as far east as Sulawesi and the Sunda Islands (see map below). While there may have been some trade between New Guinea and northern Australia in the 13th century, there is really no evidence for this, as far as I know. Thus, Australia does seem to have been a relatively ‘dark continent’ until the 17th century, especially to Europeans, and Frederick’s cockatoo does not really shed any light on that narrative.

I feel that I should emphasize how much I enjoyed the original article by Dalton and colleagues, despite my reservations above. In my view, science progresses when there is some healthy skepticism, and that’s what I have tried to present here.

Land and marine trade routes in mediaeval times.

NOTE: I accidentally posted an incomplete version of this essay a couple of days ago. My apologies for that. I blame the heat.


  • Dalton H, Salo J, Niemelä P, Örmä S (2018) Frederick II of Hohenstaufen’s Australasian Cockatoo: Symbol of Detente between East and West and Evidence of the Ayyubids’ Global Reach. Parergon 35: 35-60.

  • Frederick II (~1245) De arte venandi cum avibus. Vatican City, Biblioteca Apostolica Vaticana, codex Ms. Pal. Lat. 1071.
  • Kinzelbach R (2008) Modi auium – Die Vogelarten im Falkenbuch des Kaisers Friedrich II’. pp 62–135 in Vol 2 of Kaisers Friedrich II 1194–1250: Welt und Kultur des Mittelmeerraums (ed. Ermete K, Mamoun Fansa M, and Carsten Ritzau C). Mainz: Philipp von Zabern.
  • Rowley, I. (2018). Cockatoos (Cacatuidae). In: del Hoyo, J., Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (eds.). Handbook of the Birds of the World Alive. Lynx Edicions, Barcelona. (retrieved from on 2 July 2018).
  • Stresemann E (1975) Ornithology from Aristotle to the present. Cambridge, MA: Harvard University Press.

  • Willemsen CA (1980) Das Falkenbuch Kaiser Friedrichs II. Nach der Prachthandschrift in der Vatikanischen Bibliothek. Dortmund: Harenberg.

  • Wood CA, Fyfe FM (1943) The Art of Falconry; Being the De Arte Venandi cum Avibus of Frederick II of Hohenstaufen by Frederick the Second of Hohenstaufen. Stanford: Stanford University Press.

  • Yapp WB (1983) The illustrations of birds in the Vatican manuscript of De arte venandi cum avibus of Frederick II. Annals of Science 40: 597–634


  1. Frederick II’s manuscript: see my earlier post on this ornithologically important masterpiece here
  2. articles in popular press: but curiously not (yet) The New York Times or any of the other leading American and Canadian news media outlets.
  3. Parergon: is the journal of the Australian and New Zealand Association of Medieval and Early Modern Studies (Inc.), and has been publishing refereed articles since 1983.
  4. Frederick’s manuscript in the Vatican Library: is also available for study online in two spectacularly reproduced digital copies here and here.
  5. Quotations 1, 2, 4 & 5: from Dalton et al. 2018
  6. Quotation 3: from the headline in The Telegraph (UK)
  7. Quotation 6: Helen Dalton quoted in The Guardian (UK)
  8. previous authors on this cockatoo: see, for example, Wood and Fyfe (1943), Stresemann (1975), Willemsen (1980), Yapp (1983), Kinzelbach (2008)
  9. yellowish back and flanks: I would be inclined to interpret this colour as shading rather than as the colour of the feathers.
  10. peacocks: would probably have been a very familiar bird in the courtyards of Italy in the 13th century, having been traded along the Silk Road for centuries before. Like the cockatoo, those drawings would have been based on live speciemens
  11. likely species: Dalton et al. (2018) do seem to favour the Yellow-crested Cockatoo in their analysis, so it is curious to me that they make such a fuss about the trade routes, since the range of the Yellow-crested on one of those mediaeval routes

IMAGES: Cockatoos and peacocks from De Arte copied from the online versions; Frederick II and Silk Road map from Wikipedia; Cockatoos and range map from Handbook of Birds of the World (Rowley 2018); page from Wood and Fyfe scanned from the author’s copy.

Why Woodpeckers are Scarce in the North

BY: Bob Montgomerie, Queen’s University | 18 June 2018

On the 18th of June 1858, one hundred and sixty years ago today, Darwin claims [1] to have received that fateful letter from Alfred Russel Wallace—probably the most famous letter in the history of science. The original letter was lost but it was transcribed and read to the Linnean Society of London on 1 July and published later in their journal. That letter is well worth reading, especially because it contains some interesting insights into avian ecology. While Wallace had some useful ideas relevant to natural selection, it could be argued that his argument was not nearly as well-formed as Darwin’s [2]. In a way, his ecological and biogeographical insights are more original, in my opinion.

Wallace in 1862

Wallace wrote that letter on Ternate in the Mollucas in February 1858, sent it out on a mail steamer on 5 April. He was in the South Pacific for 8 years on a collecting trip, in part to obtain specimens that he could sell back in England but also to gather material for books that he thought, rightly so, would provide him with a lifetime of fame and fortune. He brought home more than 125,000 specimens, including more than 8000 bird skins.


I found three things to be remarkable about Wallace’s letter. First, he develops some of the same ideas about selection as Darwin,  and uses some of the same language: “state of nature”, “struggle for existence”, “stability of species”, “geometrical ratio”, “origin of…species”,  “conditions of existence”, and “superior varieties.” These are not phrases you would be likely to read in a recent paper on evolutionary biology, but may well have been argots of the scientific literature in the 1800s.

Second, he makes clear his objections to Lamarck’s ideas:

The hypothesis of Lamarck—that progressive changes in species have been produced by the attempts of animals to increase the development of their own organs, and thus modify their structure and habits—has been repeatedly and easily refuted…the view here developed renders such an hypothesis quite unnecessary, by showing that similar results must be produced by the action of principles constantly at work in nature. [3]

And third, he is remarkably insightful and creative about what today we would call evolutionary ecology with respect to passenger pigeons, woodpeckers, and clutch size.

On clutch size, he makes the perceptive observation that a species’ population size—and rate of increase—has nothing to do with the number of offspring in a brood:

…large broods are superfluous. On the average all above one become food for hawks and kites, wild cats and weasels, or perish of cold and hunger as winter comes on. This is strikingly proved by the case of particular species; for we find that their abundance in individuals bears no relation whatever to their fertility in producing offspring. Perhaps the most remarkable instance of an immense bird population is that of the passenger pigeon of the United States, which lays only one, or at most two eggs, and is said to rear generally but one young one. [3]

Great Spotted Woodpecker

On woodpeckers, he argues that they are more scarce in the temperate zone than in the tropics due to the uncertainty of overwinter food in the north, and the various morphological adaptations that would make long-distance migration difficult. I don’t even know if these observations are true, but the idea is immensely creative and demonstrates his excellent ecological insights:


Those whose organization does not permit them to migrate when their food becomes periodically scarce, can never attain a large population. This is probably the reason why woodpeckers are scarce with us, while in the tropics they are among the most abundant of solitary birds. Thus the house sparrow is more abundant than the redbreast, because its food is more constant and plentiful,- seeds of grasses being preserved during the winter, and our farm-yards and stubble-fields furnishing an almost inexhaustible supply. [3]

On the Passenger Pigeon, he reasons—correctly, I think—that its unbelievably huge populations were a product of the bird’s ability to move efficiently to track the vagaries of its occasionally superabundant food supply:

Perhaps the most remarkable instance of an immense bird population is that of the passenger pigeon of the United States, which lays only one, or at most two eggs, and is said to rear generally but one young one. Why is this bird so extraordinarily abundant, while others producing two or three times as many young are much less plentiful? The explanation is not difficult. The food most congenial to this species, and on which it thrives best, is abundantly distributed over a very extensive region, offering such difference of soil and climate, that in one part or another of the area the supply never fails. The bird is capable of a very rapid and long-continued flight, so that it can pass without fatigue over the whole of the district it inhabits, and as soon as the supply of food begins to fail in one place is able to discover a fresh feeding-ground. [3]

Like his contemporaries, however, Wallace reasoned that this species’ populations were just too big to fail: “This example strikingly shows us that the procuring a constant supply of wholesome food is almost the sole condition requisite for ensuring the rapid increase of a given species, since neither the limited fecundity, nor the unrestrained attacks of birds of prey and of man are here sufficient to check it. In no other birds are these peculiar circumstances so strikingly combined.” [3] This is one of those rare cases where we could actually learn from history and maybe not repeat Wallace’s mistake in our dealings with other species.


  • Bock WJ (2009) The Darwin-Wallace myth of 1858. Proceedings of the Zoological Society 62:1–12.

  • Davies R (2008) The Darwin conspiracy: origins of a scientific crime. London: Golden Square Books
  • Davies R (2012) How Charles Darwin received Wallace’s Ternate paper 15 days earlier than he claimed: a comment on van Wyhe and Rookmaaker (2012). Biological Journal of the Linnean Society 105:472–477.

  • Darwin CR, Wallace AR (1858) On the tendency of species to form varieties; and on the perpetuation of varieties and species by natural means of selection. Journal of the Proceedings of the Linnean Society of London, Zoology 3:46–50.

  • Davies R (2012) How Charles Darwin received Wallace’s Ternate paper 15 days earlier than he claimed: a comment on van Wyhe and Rookmaaker (2012). Biological Journal of the Linnean Society 105:472–477.

  • Gould E, Gould J, Lear E (1837) The Birds of Europe. (v. 1-5). London: pub. by the author.
  • Smith CH (2013) A further look at the 1858 Wallace–Darwin mail delivery question. Biological Journal of the Linnean Society 108:715–718.


  1.  Darwin’s claim about Wallace’s letter: Davies (2008) in particular, claimed that Darwin received the letter earlier and plagiarized it in his own notes so that he could claim priority, This seems highly unlikely to me, based on what I know of Darwin’s character and what Darwin himself says about the letter. van why and Rookmaaker (2012) present a convincing counter argument (but also see Davies 2012)
  2. Wallace’s ideas on natural selection: see Bock (2008) for details on what Wallace did have to say about selection
  3. Quotations: are all from the transcribed version of Wallace’s letter, available here

Audubon’s Legendary Experiments

BY: Bob Montgomerie, Queen’s University | 11 June 2018

In his Ornithological Biography [1], John James Audubon describes two experiments that have become legends in the annals of ornithology. Both were ingenious ideas, but Audubon’s conclusions were misleading and remained uncorrected for more than a century. According to  a new paper by Matthew Halley in Archives of Natural History, one of those experiments might never have been conducted at all.

mill grove
Audubon mansion at Mill Grove

In the spring of 1804, just before his 19th birthday [2], Audubon found an Eastern Phoebe nest in a cave near where he was living, at the plantation his father had purchased at Mill Grove, Pennsylvania [3]. Audubon had arrived from France a few months earlier and immediately began exploring the countryside where, he said “Hunting, fishing, drawing, and music occupied my every moment; cares I knew not, and cared naught about them. I purchased excellent and beautiful horses, visited all such neighbors as I found congenial spirits, and was as happy as happy could be.” [4]

Audubon wondered whether his phoebes might exhibit what we would now call ‘natal philopatry’: “I had at that period an idea that the whole of these birds were descended from the same stock.” [5]. To test that idea, he put some ‘light threads’ on the legs of the 5 nestlings. The birds, or their parents, repeatedly removed those threads but finally gave up [6]. Once the birds had habituated to the threads, Audubon says that “when they were about to leave the nest, I fixed a light silver thread to the leg of each, loose enough not to hurt the part, but so fastened that no exertions of theirs could remove it.” [7]. Others have, I think, misinterpreted Audubon’s ‘silver thread’ as being silver-coloured yarn but the term is (and was) also used for wire made of silver. Audubon later refers to these as ‘rings’ (see below) suggesting that they were solid and unlikely to be removable by the birds.


The next year, Audubon claims, he found two of those marked phoebes—which he called pewees—nesting near where they were born:

…I had ample proof afterwards that the brood of young Pewees, raised in the cave, returned the following spring, and established themselves farther up on the creek…At the season when the Pewee returns to Pennsylvania, I had the satisfaction to observe those of the cave in and about it. There again, in the very same nest, two broods were raised. I found several Pewees nests at some distance up the creek, particularly under a bridge, and several others in the adjoining meadows, attached to the inner part of sheds erected for the protection of hay and grain. Having caught several of these birds on the nest, I had the pleasure of finding that two of them had the little ring on the leg.” [8]

Now Halley—correctly I think—calls into question the veracity of these observations of natal philopatry, First, he points out, the 40% return rate (2 of 5 banded) of Audubon’s nestling phoebes far exceeds the rate of natal philopatry for this species—and indeed all passerine birds [9]—revealed in subsequent research with large sample sizes. Work done from 1988-90 at my own institution’s field station by Kelvin Conrad, for example, showed a 1.3% return rate of 217 banded nestlings [10]. A much larger study in Indiana [11] found that only 218 of 11,847 (1.8%) phoebe nestlings returned to anywhere within a 250-kmstudy area. Based on those numbers, the chance that 2 of Audubon’s 5 nestlings actually returned is almost zero.

Second, Halley checked the dates of Audubon’s return to France (12 March 1805) and subsequent arrival back in Mill Grove (4 June 1806). Based on the normal first egg dates from nearby sites, Halley argues that Audubon could not have been at Mill Grove during the 1805 breeding season and could therefore not have observed the returns he claimed. While I think Halley is correct, it is always possible that Audubon was mistaken about the years when he conducted this experiment.

My guess, though, is that Audubon did actually make up this story, not about banding the birds [12] but about them returning to their natal site. Audubon was well known to be desperate to establish his reputation as an ornithologist, in part, at least, to enhance the sales of his fabulous collection of etchings based on his paintings of North American birds. Audubon was often at odds with Alexander Wilson and George Ord, sometimes plagiarizing Wilson and fabricating evidence so that he would be seen as the first to record a species of bird or an interesting observation [13]. Elliott Coues, for example, said that Audubon: “..loved warmth, color, action; he liked to exaggerate and ’embroider,’ and make his page glow like a hummingbird’s throat, or like on of his many marvellous pictures; he had no genius for accuracy, no taste for dull, dry detail, no care for a specimen after he had drawn it.” [14]

Audubon’s other misleading experiment was designed to assess the ability of Black Vultures to detect odours, but that is a story for another day. Whether or not Audubon was always accurate in his descriptions of bird behaviour or his timelines of events, his experiments were certainly ingenious. He probably does deserve credit as the first person to band birds in North America, and he really could paint a marvellous picture:

Audubon’s Black Vultures


  • Audubon JJ (1831-1839) Ornithological biography : or an account of the habits of the birds of the United States of America; accompanied by descriptions of the objects represented in the work entitled The Birds of America, and interspersed with delineations of American scenery and manners.  5 volumes. Edinburgh: Adam & Charles Black

  • Audubon JJ (1827-1838) The Birds of America. Edinburgh & London: J. J. Audubon.

  • Audubon MR (1897) Audubon and His Journals, with Zoological and Other Notes by Elliott Coues, vol. 1. New York: Charles Scribner’s Sons.

  • Burns FL (1908) Alexander Wilson: II. The Mystery of the Small-Headed Flycatcher. The Wilson Bulletin 20:63–79.

  • Halley MR (2018) Audubon’s famous banding experiment: fact or fiction. Archives of natural history 45:118–121.

  • Weatherhead PJ, Forbes MRL (1994) Natal philopatry and the cost of dispersal in Passerine birds. Behavioural Ecology 5:426–433.

  • Weeks Jr HP (2011) Eastern Phoebe (Sayornis phoebe), version 2.0. In The Birds of North America (A. F. Poole, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA.


  1. Ornithological Biography: designed by Audubon, with the help of William MacGillivray, to accompany his paintings in Birds of North America. This 5-volume treatise (available here) contains many interesting details and insights about birds but is not an easy read.
  2. 19th birthday: Audubon was born on 26 April 1785 in the French colony that is now Haiti
  3. Audubon’s home: Audubon travelled from France to Mill Grove by himself, on the orders of his father to manage the plantation but was more interested in natural history and he eventually drove the plantation into bankruptcy
  4. quotation about hunting etc: from Audubon 1897, page ; one of the neighbours he talks about was William Blakewell, father of the woman he married a couple of years later
  5. quotation about phoebe philopatry: from Audubon (1831-39) vol 2, 1834, page 125
  6. phoebes removing threads: this thread removal and habituation must have occurred over less than two weeks as phoebes leave the nest after 16-18 days
  7. quotation about silver threads: from Audubon (1831-39) vol 2, 1834, page 125
  8. quotation about returning phoebes: from Audubon (1831-39) vol 2, 1834, page 125-127
  9. natal philopatry of passerines: Weatherhead and Forbes (1994) summarized the rates of natal philopatry for 32 migratory passerines from various published and unpublished studies. The highest recorded rate of natal philopatry was 13.5%, and the majority of species had rates <5%, much lower than Audubon recorded for his phoebes
  10. natal philopatry in Ontario: unpublished data summarized in Weatherhead and Forbes (1994)
  11. natal philopatry in Indiana: unpublished data summarized in Weeks (2011)
  12. Audubon banding birds: I am inclined to believe that Audubon did band those birds as the details ring true. He thus deserves his reputation as America’s first bander, preceding by almost a century the next attempt to band birds in the America’s
  13. Audubon’s conflicts and plagiarism: see Halley (2018) for a brief summary
  14. Elliott Coues quotation: as quoted by Burns 1908 page 68

IMAGES: Audubon mansion from Audubon (1897);  Audubon’s paintings from his Birds of America (1827-38)