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.

Ladies, Parakeets, and the Biogeography of an Extinct Bird

BY: Bob Montgomerie, Queen’s University | 25 June 2018

In 1850, an anonymous author published a superb diary of natural history observations called ‘Rural Hours by a Lady’ based on two years of exploring the woods and fields near Cooperstown, New York. The book was wildly popular, and it was not long before the author was revealed to be Susan Fenimore Cooper [1]. On page 146 she says:

Parakeet2It is well known that we have in the southern parts of the country a member of the Parrot tribe, the Carolina Parakeet. It is a handsome bird, and interesting from being the only one of its family met with in a temperate climate of the Northern Hemisphere. They are found in great numbers as far north as Virginia, on the Atlantic coast; beyond the Alleghanies, they spread themselves much farther to the northward, being frequent on the banks of the Ohio, and in the neighborhood of St. Louis. They are even found along the Illinois, nearly as far north as the shores of Lake Michigan. They fly in flocks, noisy and restless, like all their brethren…In the Southern States their flesh is eaten…Birds are frequently carried about against their will by gales of wind; the Stormy Petrels, for instance, thoroughly aquatic as they are, have been found, occasionally, far inland. And in the same way we must account for the visit of the Parakeets to the worthy Knickerbockers about Albany.  [2]

Here, she correctly describes the bird as being most common in the southeastern states, though seen regularly as far north as the Great Lakes west of the Allegheny Mountains. What she did not know, of course, was that these were two subspecies, with different morphologies, ecologies and migratory strategies, as described below.

Shufeldt’s photo

The Carolina Parakeet was still abundant throughout its range in 1850 but, like the Passenger Pigeon, was soon to be extirpated. The second-last individual was a female called ‘Lady Jane’ who died at the Cincinnati Zoo in 1917; the last of its kind being Lady Jane’s mate, a male named ‘Incas’ who died there in 1918, one hundred years ago. Coincidentally, Incas died in the same cage where Martha, the last passenger pigeon, had died in 1914 [3]. There were reports of sightings in the wild for another 40 years or so, in Florida and Georgia, but none of those records were authenticated. Among the North American birds that have become extinct since the arrival of Europeans, the biology of the breeding biology Carolina Parakeet may be the poorest known [4]. And there is, surprisingly, only one photo of the bird in a natural-looking setting [5], taken by the irrepressible Robert Shufeldt in about 1900.


A recent pair of papers by Kevin Burgio and colleagues uses all of the known specimens and sightings of this bird to reveal some interesting insights into its distribution, ecology, and taxonomy. There were 401 of those sightings recorded between 1564 and 1944, and nearly 800 specimens in museums and private collections worldwide [6], almost all collected in the 1800s. As shown on the graphs below, the number of records climbed exponentially from 1500 to 1900, reflecting the increases in exploring the new continent, in writing about natural history, and in preserving ornithological data and specimens. There was an uptick in collecting, or at least preserving specimens, from 1870-1900 when it became clear that the bird was disappearing [6].

Records and specimens with known dates—note the log scale on upper two graphs.

Analyzing records only from states where the parakeet was known to breed, Burgio and colleagues, georeferenced all the data and used 147 unique localities to create the species breeding distribution models shown on the map to the right below. The map on the left was produced in 1889 by Edwin Hasbrouck with the known range in his day (black shading) nicely matching the newly reconstructed ranges of the two subspecies.

Burgio and colleagues’ research also suggested (i) that the breeding range of this species was much smaller than previously thought, (ii) that the two subspecies, previously only vaguely defined by size and colour, actually had disjunct ranges and occupied somewhat different climatic niches, and (iii) that the western subspecies was almost certainly migratory where the eastern one was not.

The authors also hoped their analysis would help to inform current conservation practices in an effort to save the 8% of bird species currently threatened to disappear as a result of climate change. Parrots, in particular, are in bad shape, with 42% of species listed as threatened or endangered.

LEFT from Hasbrouck (1889) estimating the limits of the parakeet’s historical range (black line) with shading showing the range in the 1880s. RIGHT from Burgio et al. (2017) estimating the ancestral breeding ranges of the two subspecies (Hasbrouck’s range limit shown as a red line)


Clay pipe ca. 1650

As much as I like those recent papers, I think it’s unfortunate that many biogeographers draw their maps as if animals obeyed political boundaries, as on the right-hand but not the left-hand maps above. The right-hand graph implies, for example, that the bird could never have crossed the US-Canada border as there was nowhere to go. Despite that, there is some evidence that it did occasionally occur in southwestern Ontario, possibly blown off course as Susan Cooper suggested above. At an archaeological dig at Grimsby, Ontario, for example, Walter Kenyon found a clay pipe that looks distinctly like a parrot, made by native peoples in the mid-1600s. And Rosemary Prevec found 3 Carolina Parakeet bones at a native site near London, Ontario, dated at around 1100 CE. Both of these findings are no more than suggestive and could have been obtained in trade with natives living further south.

Possibly more convincing are some observations that Samuel de Champlain recorded in his notes in 1615, in the woods near where I live in Kingston. He says that he  “…penetrated so far into the woods in pursuit of a certain bird which seemed to be peculiar, with a beak almost like that of a parrot, as big as a hen, yellow all over, except for its red head and blue wings, which made successive flights like a partridge.” [8] There are definitely no other birds even remotely resembling that description in eastern Ontario today.

Catalog page, New York Millinery and Supply Company, Inc., New York

None of this nationalism is really important to our understanding of the bird’s ecology and demise, except to note that at one time the species was clearly widespread and mobile. What is important is an attempt to understand why they went extinct, as even by the middle of the 1800s it appeared to be declining in numbers [6].

Burgio and colleagues point to habitat destruction and hunting as the likely causes. Not surprisingly, the parakeet’s feathers were prized for the millinery trade, with some reports suggesting that ladies hats were sometimes adorned with skins of the whole bird. The 1901 ad to the right, for example, shows a whole parrot (skin) in the lower right corner, for the bargain price of 25¢ a bird or $2.95 a dozen (about $7.50 and $88 in today’s currency). While the documentation is sketchy, it is also likely that this species was a popular cage bird in Germany as well as in North America. The only other known photo, besides Shufeltdt’s, is also one of a pet called  ‘Doodles‘, kept by Smithsonian malacologist Paul Bartsch. In 1900, ‘doodle‘ meant ‘fool‘ and not the ‘absentminded scribble‘, Google commemorative, or online scheduler that it is today. And I wonder if Bratsch gave it that name to reminder him what fools we are when let any species go extinct.


  • Anonymous [Cooper, SF] (1850) Rural Hours by a Lady. New York: G. Putnam.
  • Burgio KR, Carlson CJ, Tingley MW (2017) Lazarus ecology: Recovering the distribution and migratory patterns of the extinct Carolina parakeet. Ecology and Evolution 7:5467–5475.
  • Burgio K, Carlson C, Bond A (2018) Georeferenced sighting and specimen occurrence data of the extinct Carolina Parakeet (Conuropsis carolinensis) from 1564-1944. Biodiversity Data Journal 6:e25280.
  • Cokinos C (2000) Hope Is the Thing With Feathers: A Personal Chronicle of Vanished Birds. New York: Penguin.
  • Fuller E (2013) Lost Animals: Extinction and the Photographic Record. London: Bloomsbury.

  • Greene WT, Dutton FGFG, Fawcett B, Lydon AF (1883) Parrots in Captivity, v. 2. London: George Bell and Sons.

  • Hahn P (1963) Where is that Vanished Bird? Toronto: University of Toronto Press. [see this previous post for more on this book[
  • Kennedy CC (1984) Did Champlain stalk a Carolina Parakeet in southern Ontario in 1615? Arch Notes 84:55–62.

  • McKinley, D. (1960) The Carolina parakeet in pioneer Missouri. The Wilson Bulletin 72:274–287.
  • McKinley D (1977) Climatic relations, seasonal mobility, and hibernation in the Carolina Parakeet. Jack-Pine Warbler 55:107–124.
  • Prevec R (1984) The Carolina Parakeet—its first appearance in southern Ontario. Arch Notes 84:51-54.
  • Snyder NFR (2004) The Carolina Parakeet: Glimpses of a Vanished Bird. Princeton, NJ: Princeton University Press.


  1. Susan Fenimore Cooper: was a superb naturalist, author, and artist whose work was overshadowed in more ways than one by that of her father, James. She deserves recognition and a separate essay on her own work, Stay tuned.
  2. Cooper quotation: from Anonymous 1850 page 146
  3. ‘Incas’ the parakeet: Like Martha, the last Passenger Pigeon, Incas was frozen and sent to the Smithsonian, but he was lost in transit (Fuller 2013)
  4. breeding biology poorly known: see Snyder (2004)
  5. Shufeldt’s photo: is one of a pair of pet birds that Shufeldt borrowed from his friend Edward Schmidt, and it took him hours to get it to sit still enough on a cocklebur to make a decent photo (Cokinos 2001). Both of Schmidt’s birds later died from chewing on the bars of their cage, possibly from lead paint poisoning (Fuller 2013)
  6. declining numbers by mid 1800s: see Hasbrouck (1889)
  7. records and specimens: see Hahn (1963), McKinley (1960, 1977) and Snyder (2004) for background
  8. Champlain quotation: from Kennedy 1984 page 55

IMAGES:  first parakeet is by Robert Ridgway from Baird et al. (1905); graphs by the author based on data in Burgio et al. (2017, supplement)—parakeet is an engraving by Benjamin Fawcett in Greene et al. (1883); maps from the original papers; clay pipe from Kennedy (1984); millinery ad from the Smithsonian collection

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)

A Whiskey-Jack by Any Other Name

BY: Bob Montgomerie, Queen’s University | 4 June 2018

Canada Jay, Algonquin Park, 1968

The year I turned 21, I got my dream job: seasonal naturalist at Algonquin Provincial Park. Algonquin, established in 1893, was only the second provincial park to be created in Canada, and the first to be designated to protect a natural environment [1].  This vast ‘wilderness’ area (7653 km2) is only 3 hours by car from Ottawa, 4 from Toronto, and 5 from Montreal. The park now gets more than a million visitors a year, concentrated mainly along the highway that runs east-west across its southern edge. At the time, I had birded and bird-banded only in southwestern Ontario, so this was a chance to see some boreal species on their breeding grounds, species that I had only seen previously on migration or in winter, if at all—Common Loons, White-throated Sparrows, Saw-whet Owls, Pine Grosbeaks, Red Crossbills, Ravens, Three-toed Woodpeckers, and Gray Canada Jays.

Because I was no longer in school, I started work at Algonquin two months earlier than the other half dozen seasonal naturalists, mainly to get the museum’s [2] specimen collections in order and to prepare for the onslaught of summer visitors. Seasonal naturalists were hired to interact with the visitors who flooded the park in July and August during the public schools’ summer vacation. I arrived at Algonquin at the end of April where I met Russell J. Rutter, the only full-time park naturalist, who lived near the small town of Huntsville, a half hour west of the park. Rutter was a crusty old guy but we got on well and he often took me out birding, botanizing, hiking the trails where we would lead nature walks, and howling for wolves [3].

One day, as we walked along some abandoned railway tracks, 3 Canada Jays appeared at the edge of the woods. Russ made a whispery-squeaky sound and all three birds flew right up to us, one landing on Russ’s hand to get some food that he had brought with him just for that purpose. Two of those jays had colour-bands (WR, and YORL [4]), and this was a family group, Russ said. Russ told me he had decided a few years earlier to start studying Canada Jays so that he could follow individuals through their lives. He had already discovered that they often travel in family groups, hoard food for the winter, and occupy year-round territories. I was enthralled—I had no idea that the birds I had watched breeding in southern Ontario were not doing what all birds did.

Russ was also furious with the AOU for changing the bird’s common name from Canada Jay to Gray Jay in its 1957 checklist. I must admit I was not really even aware of this as my field guide—the 3rd edition of Peterson, published in 1947—called them Canada Jays and that’s what I would have called them but I had never seen one before. That summer, we naturalists often told the park visitors about the name change but also that the Native Algonquins had once called them whisky-jacks, which made for a good story.

from the AOU Checklist 1957

One of the seasonal naturalists that year was Dan Strickland, who had been an Algonquin Park naturalist the previous summer or two and was now studying Canada Jays (‘Geais Gris’) for his MSc thesis at the Université de Montréal. Dan was conducting his research in la réserve faunique de La Vérendrye in Québec, inspired by Rutter’s work in Algonquin. Dan’s focus was on their social behaviours and their food-caching strategies. He stayed at Algonquin as Park Naturalist for the rest of his career, and has continued to work on the Canada Jay for the past half century, one of the longest continuous studies of a bird species worldwide. Not surprisingly, Strickland was instrumental in the recent official change of the species’ name back to Canada Jay just last month. The popular press in Canada is all abuzz about this change, in part because calling them ‘Canada Jays’ may enhance the possibility that this species will now be recognized as Canada’s national bird.

Martinet’s etching of Canada Jay in Brisson (1760)

The Canada Jay was first described for science in 1760 as ‘Le Geay brun de Canada’ (Garrulus Canadensis fuscus) by the French zoologist  Mathurin Jacques Brisson in his Ornthologie, with a remarkably good illustration by François Nicolas Martinet. An English translation of Brisson’s common name might be the ‘Canada Brown Jay’ [5]. This description, and the illustration, were based on a specimen in the collection of René Antoine Ferchault de Réaumur, as neither Brisson nor Martinet had seen the bird in the wild. de Réaumur was primarily an entomologist but he had an extensive  private museum where he employed Brisson as what we would today call a ‘curator’.  Brisson’s description was the basis for Linnaeus presenting the bird’s  ‘official’ scientific name as Corvus canadensis, in 1766, but Linnaeus did not suggest a common name, as that was not the purpose of his work.

Thomas Pennant was probably the first to publish an English common name—Cinereous Crow [6]— for this species, in his Arctic Zoology of 1784. Pennant had never seen the bird alive either, relying instead on Samuel Hearne’s description [7]. Hearne joined the British Navy at the age of 11 and then the Hudson’s Bay Company at 21, stationed first at Fort Prince of Wales (now Churchill) in Manitoba. While there, he made 3 major expeditions into the interior of present-day Nunavut, in 1771 reaching the mouth of the Coppermine River where it drains into the Arctic Ocean. Hearne’s Journey, published posthumously in 1795, contains detailed species accounts of at least 50 bird species that he encountered on his expeditions, with remarkable insights into their behaviours and ecologies. Hearne called this jay the ‘Cinereous Crow (Perisoreus canadensis)’, noting in his account that “ is called by the Southern Indians [presumably Cree], Whisk-e-jonish, by the English Whiskey-Jack, and by the Northern Indians [presumably Chipewyan] Gee-za…” [8]

Then, in 1829, John Richardson called this bird ‘The Whiskey-Jack (Garrulus canadensis)’ in his comprehensive Fauna Boreali-Americana, coauthored with William Swainson. Richardson had explored northern Canada with the Franklin Expeditions of 1819-22 and 1825-27, and would have had first hand experience with this species. A half-century later, with the publication of the first AOU checklist in 1886, the official names became Canada Jay and Perisoreus canadensis, and would remain that way until 1957.

from the AOU checklist 1886

During the Washington AOU conference in 2016, Dan Strickland mined the AOU archives at the Smithsonian to figure out how and why the common name of this species was changed by the AOU in 1957. Based on that information, he made a very compelling case to have the name changed back to Canada Jay. The details are complex but nicely outlined by Strickland here and here.

While the scientific names of birds (and all plants and animals) are assigned following some strict rules, there are no such rules for common names. Attempts to make some rules for common names have not been successful (see here for example), and for good reason, I think. Common names say something—not always useful (see here)—about appearance, vocalizations, habitat, and localities, as well honouring people—not always logically (see here)—and their contributions to science or discovery. Common names are what most of us use when we talk about birds and ‘official’ common names should probably, like all language, reflect usage rather than some esoteric rules. No amount of rule-making will stop duck hunters from using the names ‘whistler’, ‘sprig’, ‘greenhead’, ‘bluebill’, spoonbill’ or ‘skunkhead coot’ [9], nor the visitors to Algonquin Park from calling their favourite bird the ‘whiskey-jack’.


  • American Ornithologists’ Union (1886) The code of nomenclature and check-list of North American birds. New York: American Ornithologists’ Union.
  • American Ornithologists’ Union (1957) Check-list of North American birds. Ithaca, N.Y.: American Ornithologists’ Union.
  • Brisson M-J, Martinet FN (1760) Ornithologie, ou, Méthode contenant la division des oiseaux en ordres, sections, genres, especes & leurs variétés (t.2 ). Parisiis: Ad Ripam Augustinorum, apud Cl. Joannem-Baptistam Bauche, bibliopolam, ad Insigne S. Genovesae, & S. Joannis in Deserto.
  • Hearne S (1795) A journey from Prince of Wales’s Fort in Hudson’s Bay to the Northern Ocean. London: Strahan and Cadell.
  • Houston CS, Ball T, Houston M (2003) Eighteenth-century Naturalists of Hudson Bay. Montreal: McGill-Queen’s University Press.
  • 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)), 12th edition. Holmiae :Impensis direct. Laurentii Salvii.
  • Pennant T (1784-85) Arctic Zoology, 2 vols. London: Henry Hughs
  • Rutter, R.J. 1969. A contribution to the biology of the Gray jay (Perisoreus canadensis). Canadian Field-Naturalist 83: 300-316.
  • Strickland D (1969) Écologie, comportement social et nidification du Geai Gris (Perisoreus canadensis). Master’s Thesis, Univ. Montréal, Montréal.
  • Strickland D (2017) How the Canada Jay lost its name and why it matters. Ontario Birds 35: 2-16.
  • Swainson W, Richardson J (1831) Fauna boreali-americana, or, The zoology of the northern parts of British America: containing descriptions of the objects of natural history collected on the late northern land expeditions, under command of Captain Sir John Franklin, R.N. Part second, the birds. London: John Murray.


  1. Canadian provincial parks: Queen Victoria Park at Niagara Falls was Canada’ first provincial park, established in 1885 in part to clean up the area around the falls and reduce the incidence of crime
  2. Algonquin park museum: in those days was on the shore of Found Lake but now is a spectacular Visitor Centre 30 km east of the old museum
  3. howling for wolves: the Algonquin Wolf Howl, held in August every year, attracts more than a thousand visitors who gather at the roadside at sunset, hoping to hear the wolves respond to the park naturalists’ attempts to imitate the howls. Those howl imitations are so good that one year the naturalists mistakenly called back and forth to each other, each group assuming that they were hearing real wolves.
  4. colour-bands: the letters refer to the colours, usually left-leg top and bottom, then right-leg top and bottom. So these birds were white on the left, red on the right, and yellow over orange on the left and red over light-blue on the right. Researchers like band combos that they can pronounce so these birds were ‘whir’ and ‘yorel’ to Rutter.
  5. Canada Brown Jay: in Brisson’s day, ‘Canada’ referred to the French colony that we now call Québec.
  6. Cinereous Crow: ‘cinereous’ means ‘ashy-grey’ which correctly describes the bird’s colour but we can be grateful that this name was never used after the middle of the 19th century. I have little doubt that early explorers to Canada in the 16th and 17th centuries will have said something about this species but I cannot find an earlier reference than Brisson (1760)
  7. Samuel Hearne’s description: we know that Hearne gave Pennant a copy of his observations while Hearne was in England during the winter of 1782-83 (Houston et al. 2003)
  8. quotation about Native names: from Hearne (1795, page 374)
  9. hunter’s names for ducks: officially (AOU checklist) these are Common Goldeneye, Norther Pintail, Mallard, Lesser (or Greater) Scaup, Northern Shoveler, and Surf Scoter, respectively

IMAGES: Canada Jay photo by the author; pages from AOU checklists and Brisson are in the public domain

The Spring Rivalry of Birds

BY: Bob Montgomerie, Queen’s University | 28 May 2018

Unless you have been living in a cave for the past couple of months, you will have been well aware of the spectacle of mature, powerful males posturing to one another, showing off their weapons, advertising their prowess, and intimidating their neighbours. Some of them even coerced females into unwanted sexual acts, or spent some time cheating on their mates. I refer, of course, to what Charles Bethune Moffat called ‘The Spring Rivalry of Birds‘ in his 1903 paper in The Irish Naturalist.

American Coots fighting at a territory boundary (PHOTO Bruce Lyon)

Moffat was born in January 1859 on the Isle of Man in the Irish Sea, 10 months before the publication of Darwin’s Origin of Species. Moffat’s family soon moved to Ireland where he grew up and eventually spent his entire professional career as a lawyer and journalist.

IrishNatBut Moffat was also a consummate naturalist who published about birds in popular articles in Dublin’s Daily Express, and in scientific papers in The Irish Naturalist [1] and The Irish Naturalists’s Journal. His paper on the ‘spring rivalry’ is best known because it was probably the first publication to present, in English [2], the idea that male birds defend a territory: “…that the battles fought between the male birds in spring have for their object, not the winning of particular females, but the acquiring of suitable plots of land, and that the song and bright plumage of the male are a warning to other males and an intimation to a female that a suitable territory has been acquired.” [3]

Reading Moffat’s spring rivalry paper, I get the distinct impression of a man who was thoughtful, logical and a keen observer—all traits that would have made him an excellent lawyer and journalist (both of which he was) and a really good scientist. He noted, for example, “…that we seldom find in close proximity to each other two nests belonging to the same species of bird.”, “…that cock birds in early spring spend a great deal of their time in fighting one another.”, and “..that we must have a very large number of non-breeding birds of both sexes, prevented from breeding simply by the fact that they have no suitable ground.” [3]

Moffat liked Darwin’s ideas about evolution and natural selection but he thought—like many others at the time—that Darwin was simply wrong about sexual selection, especially with regard to female choice:

…the number of hen-birds, so far as we can gather, is fully equal to the number of cock-birds, so that when all the fighting is over, there is nothing to prevent all the birds from marrying and settling down to ‘ live happily ever after.’ We may suppose, in our sentimental way of looking at things, that even then the poor beaten cock-bird suffers from a certain amount of depression when he thinks of the greater charms of her to whom he first paid court. But we have the assurance of experts that no such thing happens; that one hen-bird is quite as good as another, and that every cock-bird is perfectly content with the first mate he can get. That is sometimes laid down as the reason why hen-birds, as a rule, have not developed bright nuptial colours or melodious voice…I think we must infer that there is very little free choice or aesthetic selection, and that the hen bird is mainly guided by prudential motives in accepting the owner of the soil. [3]

Dark-eyed Junco fighting his reflection in a car window (PHOTO: Bruce Lyon)

Having observed male Blackbirds and Chaffinches continually battling their reflections in his windows, Moffat quite perceptively concluded that these fights were about space, as there was no female in sight and those reflected interlopers had nether sung nor shown any interest in the male’s mate: “the imaginary enemies, on whom so much fury was expended, were guilty of no crime beyond that of being in the spot where they were.” [3]. Even when 35 Starlings were shot one at a time, consecutively, from a single breeding pair, the remaining bird almost immediately got a new partner, suggesting to Moffat that birds did not really care who they mated with, and that space was therefore of primary importance.

But why ‘space’? Moffat was also skeptical about Darwin’s idea that—at least in birds—many more offspring are produced than could possibly survive to reproduce: “I cannot believe that the theory of Natural Selection — for which I have a great respect, and which I must carefully guard myself against appearing for a moment to call in question— requires this sacrifice, or anything like it…as regards birds I am altogether unable to find grounds for believing in so great a death-rate..” [3]. Indeed, his censuses of his local House Martins suggested to him that the annual mortality rate was very low, for just about as many birds arrived back in spring as had departed the previous autumn [4].

VCWE1962His solution to this conundrum of territoriality—anticipating V. C. Wynne-Edwards’s ideas on group selection published 60 years later—was: “..that there are checks of a prudential kind on the marriage of birds, and that these checks may be a very important factor in keeping the number of birds absolutely permanent.” [3]. Thus Moffat argued that birds were simply being prudent, dividing up the available land in such a way that would keep the population stable, without the needless waste of reproduction wherein the excess individuals would die off as Darwin had argued.

He was certainly not alone in his skepticism about sexual selection and population regulation, and it took the reaction to Wynne-Edwards’s ideas—mainly by David Lack and George Williams—to set the record straight with regard to the logic of natural selection. And that set the stage for a re-examination of sexual selection and its influence on the songs, plumage colours and ornaments, and courtship behaviours of birds—traits that benefit individuals not species.

So, next time you hear the twitter of North America’s Orange-crowned Warbler, see European Coots threaten their neighbours, watch the Siberian Grouse perform his flutter-jump, or hear the Oriental Cuckoo whistle his ridiculous ‘poo-poo’, remember that none of this male behaviour is performed for the good of the species.


  • Altum B (1868) Der Vogel und sein Leben. Münster: Niemann. [available here]

  • Kennedy PG (1946) Charles Bethune Moffat. (1859-1945). British Birds 39:81–82.

  • Moffat CB (1903) The spring rivalry of birds. Some views on the limit to multiplication. The Irish Naturalist 12:152–166.

  • Stresemann E (1947) Baron von Pernau, pioneer student of bird behavior. The Auk 64:35–52.

  • von Pernauer FJA [published anonymously without his consent] (1702) Unterricht, was mit dem lieblichen Geschöpff, denen Vögeln, auch ausser dem Fang, nur durch die Ergründung deren Eigenschafften und Zahmmachung oder anderer Abrichtung, man sich vor Lust und Zeitvertreib machen könne, gestellt Durch den Hoch- und Wohlgeborhrnen Hn. Freyherrnn: Herrn von P [ available here]
  • White G (1772) Letter XI to The Honourable Daines Barrington. [available here]

  • Wynne-Edwards VC (1962) Animal Dispersion in Relation to Social Behaviour. Edinburgh: Oliver and Boyd.


  1. papers in Irish Naturalist: he published 40 papers and short notes in this journal, as well as 40 book reviews
  2. first presentation in English: Ferdinand Johann Adam von Pernauer (1702) and, especially, Bernard Altum (1868) had previously introduced and developed the idea of territoriality and the concept of ‘territory’ in birds. Moffat also notes that Gilbert White had a similar view that he expressed in a letter written in 1772: “such a jealousy prevails between the male birds that they can hardly bear to be together in the same hedge or field…it is to thisspirit of jealousy that I chiefly attribute the equal dispersion of birds in the spring over the face of the country.”
  3. quotations: all from Moffat (1903)
  4. House Martin observations: his mistake here is that he assumed that these were the same individuals, not realizing that overwinter mortality would have left openings to be filled by birds that might otherwise have been non-breeders

George Ord’s warbler

BY: Bob Montgomerie, Queen’s University | 21 May 2018

Cape May Warbler (Wilson 1812, part of Plate LIV)

I have always liked the Cape May Warbler. The male in spring is a handsome bird, but scarce enough here in eastern Ontario that I see only one or two every spring [1]. When I first saw one on migration at Long Point Bird Observatory in the 1960s, my friend and mentor David Hussell said that it was really a misnomer—the species was originally collected on Cape May, NJ, he said, but had hardly ever been seen there since.

There are only three other North American birds named after such a small region of the continent [2], and the others are also misnomers in the sense that they do not represent anything useful about the bird’s range, habitat, appearance or song [3]. While there are rules about priority in the assigning of scientific names to species, common names are largely at the whim of the first person to describe the species, and, in North America, the later machinations of the AOU Check-list committee (now AOS’s North American Classification Committee).

That ‘first’ Cape May Warbler was a male collected by George Ord on a collecting trip with Alexander Wilson on Cape May in May 1812 [4]:

THIS new and beautiful little species was discovered in a maple swamp, in Cape May county, not far from the coast, by Mr. George Ord of this city, who accompanied me on a shooting excursion to that quarter in the month of May last…The same swamp that furnished us with this elegant little stranger, and indeed several miles around it, were ransacked by us both for another specimen of the same; but without success. Fortunately it proved to be a male, and being in excellent plumage, enabled me to preserve a faithful portrait of the original. [5]

Wilson described the bird, painted it (see above), and called it the Cape May Warbler Sylvia maritima in Volume 6 (page 99) of his American Ornithology, published in 1812. It was not seen again on Cape May until September 1920 when Witmer Stone found it there but it has been increasingly recorded—and is now regularly seen—on Cape May during migration ever since [6].

Unbeknownst to Wilson, the bird had already been described  as Motacilla tigrina in 1789 in Johann Friedrich Gmelin’s edition of Linaeus’s Systema naturae:

top of page 985 in Gmelin (1789) describing Motacilla tigrina

But wait, Gmelin says that this is the same species as Brisson’s Ficedula canadensis fusca and F. dominicensis, Buffon’s ‘Figuier tacheté de jaune,’ Edwards’s ‘Spotted Yellow Flycatcher,’ and Pennant’s (1874) ’Spotted Yellow Warbler.’ OK, Buffon, Edwards and Pennant did not give it a scientific name, and Brisson’s F. canadensis is the Chestnut-sided Warbler and his F. dominicensis is the Mangrove Warbler. So Gmelin is considered to be the scientific naming authority for this species, and the species name he gave it, tigrina, is official. He also correctly identified the habitat as ‘Canada’, presumably based on Pennant who said “Inhabits also Canada, which may be its place of summer residence and breeding.” [7]

But how did Gmelin (and the others) who wrote in 1789 even know about this species if the ‘first’ specimen was the one collected by George Ord on Cape May in 1812? The short answer is that Wilson was mistaken, as was Charles Lucien Bonaparte who later wrote about Wilson’s nomenclature [8] and edited editions of Wilson’s American Ornithology after Wilson died in 1813. Since Gmelin, Brisson, Buffon and Pennant were all referring to Edwards’s specimens, Gorge Ord’s warbler was actually the third specimen of what we now call the Cape May Warbler.

Edward’s (1758) Spotted Yellow Fly-catcher (L) and Yellow-tailed Flycatcher (R)

The first specimen of what we now call the Cape May Warbler was actually collected on 1 November 1751 when a male and female landed on a boat about 56 km off the coast of the island of Hispaniola in the Caribbean. George Edwards obtained the specimens and published his painting of the male in 1758 on a plate with an American Redstart (shown to the right). In the plate caption [9], Edwards says these “undescribed small Birds” were collected by Thomas Stack “in a voyage from London to Jamaica.” Of the  female, Edwards says: “the breast in the hen was of a dirty yellow white spotted with dusky, and something less bright on the back; otherwise they are marked very much alike. These birds I believe have never been figured or described until now.” [10]

I have actually always liked the name ‘Cape May Warbler.’ Even though that moniker does not tell us anything useful about the bird, the name has an interesting history, in the same way as the name ‘Lady Ross’s Turaco’ that I wrote about in a previous post. And it’s much better than Boat-off-the-coast-of-Hispaniola Warbler, or Spotted Yellow Fly-catcher.


  • Bonaparte CL (1826) Observations on the Nomenclature of Wilson’s Ornithology. Philadelphia: Anthony Finley.
  • Brisson M-J, Martinet FN (1760) Ornithologie, ou, Méthode contenant la division des oiseaux en ordres, sections, genres, especes & leurs variétés (t.1 (1760)). Parisiis :Ad Ripam Augustinorum, apud Cl. Joannem-Baptistam Bauche, bibliopolam, ad Insigne S. Genovesae, & S. Joannis in Deserto.
  • Burtt Jr EH, Davis Jr WE (2013) Alexander Wilson: The Scot Who Founded American Ornithology. Cambridge, Mass.: Harvard University Press.
  • Edwards, G (1758) Gleanings of Natural History. Part I. London: Printed for author at the Royal College of Physicians.
  • Gmelin JF (1789) Caroli a Linné. Systema naturae per regna tria naturae : secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. (Tom. 1 Pars. 2). Lipsiae: impensis Georg. Emanuel. Beer.
  • Pennant T (1784) Arctic Zoology, 2 vols. London: Henry Hughs.
  • Wilson A (1908-1914) American Ornithology; or, the natural history of the birds of the United States. Vols I-IX. Philadelphia: Bradford and Inskeep. Available here.


  1. Rarity of Cape May Warbler in eastern Ontario: ebird says I am not alone as almost all sightings within 100 km of where I live in Kingston, Ontario, are of 1 bird on a given day.
  2. Area of Cape May: depending on how you measure it, Cape May is no more than 50 km2
  3. Three other North American bird misnomers: Nashville Warbler also named by Wilson in 1811 based on a specimen he collected on migration near Nashville—the bird breeds in the boreal forest of eastern Canada and northeastern USA, as well as in the mountains from BC to California; Philadelphia Vireo named in 1851 by John Cassin based on a bird collected on migration in Bingham’s Woods near Philadelphia where he lived and worked—the bird breeds mainly in Canada from British Columbia to Newfoundland; and the Key West Quail-dove which did once breed in the Florida Keys (including Key West) but is now just a vagrant in Florida having been extirpated as a breeder there in the 1800s—the bird was described by Charles Bonaparte in 1855 based on a specimen from Key West but that was even then the very northern tip of its breeding range that encompasses the Greater Antilles and the Bahamas.
  4. Collected in 1812: since Wilson was writing in the summer of 1812, Burtt and Davies (2013) interpret ‘May last’ as meaning May 1812, but Ord later claimed that the bird was shot in May 1811, possibly as an attempt to show that his bird was collected before a bird that Audubon thought might be the same species. Audubon collected his bird in Kentucky in May 1811, and called it the Carbonated Warbler. See Burtt and Davies (2013 page 341) for further information on this.
  5. Quotation from Wilson: 1812 (vol 6), page 99
  6. Sightings on Cape May: ebird, for example, shows more than 100 sightings since 2011.
  7. Quotation from Pennant: 1784 page 407. Pennant actually called it the Spotted Warbler and not the Spotted Yellow Warbler as Gmelin claimed.
  8. Bonaparte’s nomenclature: see Bonaparte 1824, page 14
  9. Edwards’s plate: is numbered ‘257 and appears between pages 100 and 101 in Edwards (1758)
  10. Quotation from Edwards: 1758 page 102

IMAGES: all from digital copies of the books in the Biodiversity Heritage Library, in the public domain

Lumpers and Splitters

BY: Bob Montgomerie, Queen’s University | 14 May 2018

Almost everyone I know who is interested in birds has their favourite group, and the reasons for those favourites vary widely. For some, the difficulty—and their mastery—of field identification is most appealing: I am thinking peeps, LBJ sparrows, fall wood warblers in North America and leaf (Phyloscopus) warblers in Europe. For others, it’s the beauty of—variously—their songs (wood warblers, thrushes, mimics), their plumages (male wood warblers in spring, hummingbirds, trogons, peafowl), their ability to fly (falcons, hummingbirds, swallows, swifts), their intelligence (corvids, tits, parrots), or their way of life (seabirds, hummingbirds, waterfowl, shorebirds, antbirds, brood parasites, cooperative breeders, lekking species, owls).

Peeps at Moss Landing, California

For many scientists, their favourites are the birds that they have studied most—often as much by accident as by design. Among my ornithologist friends and colleagues, the favourites are coots, murres, manakins, Parids, fairy wrens, acorn woodpeckers, barn swallows, anis, and snow geese. My own favourites are hummingbirds (PhD research), shorebirds and ptarmigan (20 years of arctic research), and seabirds (summer job before PhD).

When I began birding, however, my favourites were the Hylocichla (now Catharus) thrushes and the Empidonax flycatchers, possibly because some species were hard to identify. I was also intrigued by the geographic mosaic of Empidonax ranges and the incredible similarities in their plumages described by Ned Johnson, among others. Why is it that some species in some genera (Dendroica males in springfor example) are so easy to tell apart whereas in other genera (like Empidonax) species can only be distinguished morphologically by careful measurements of a bird in the hand (or so we thought). My own careful examination of Empidonax flycatchers while banding them at Long Point Bird Observatory resulted my first paper in an international journal more than 50 years ago.

Willow (L) and Alder (R) Flycatchers

Because of that early interest in the Empidonax flycatchers, I was both dismayed and intrigued when the AOU split the Traill’s Flycatcher into two species—Alder and Willow—in 1973 on the basis of their song and some morphological traits. Here was a species that I thought I knew well (albeit only on migration), that was actually two species, even more cryptic than the other hard-to-tell-apart species in that genus. This was my first lesson in the fluidity of taxonomy and the possibility that there were almost certainly other cryptic species to be found. I remember wondering, for example, if the Grey-cheeked Thrush might be two species, based on the obvious morphological and geographic differences in the subspecies bicknelli, and that, too, was formally recognized as a separate species by the AOU in 1995.

Throughout the 20th century, there was often a tension between lumpers and splitters [1] when it came to avian taxonomy. Some suggested that the differences between splitting and lumping were like the differences between liberal and conservative governments in democracies. We now recognize, however, that periods of lumping and splitting are the result of changing opinions about species definitions, and the progress of technologies that allow us to distinguish among populations.

A new paper in PLoS ONE by Gaurav Vaidya, Denis Lepage, and Robert Guralnick [2] uses the 136 years of AOU checklists to examine, among other things, the historical patterns of lumping and splitting of the North American avifauna by taxonomists. The first AOU checklist was published in 1886, the production of which was one of the stated reasons for the for the formation of the American Ornithologists’ Union in 1883. Since then, 6 additional full ‘editions’ of the checklist have been published, with 57 supplements published at roughly 2-year intervals between editions.

Since 1886, fully three-quarters of the ~900 North American species have not been involved in what the authors call the ‘correction process’, being either lumped with other species or split into two or more species by the checklist committees. Sixteen percent have been corrected once, and the remaining 10% twice or more. In all, there have been more  lumps (142) than splits (95) in the past 130 years but the pattern of change has been striking. Lumping was initially the norm, probably largely due to the widespread adoption of the Biological Species Concept. But very few species have been lumped since 1980 as the rate of splitting took off, likely because DNA technologies improved and there began a (slight) shift toward a phylogenetic species concept.


Their analysis identified three periods of relative stasis, when there were no corrections to be made, perhaps roughly coinciding with periods of societal upheaval and dramatic scientific changes in taxonomic philosophies and practices.

Vaiyda and colleagues also identify some intriguing examples of  “… a current, ongoing taxonomic recorrection process, in which corrections made in the first half of the 20th century are now being reverted in light of new evidence and better tools. [3]. To the non-scientist this must look like the taxonomists are incredibly indecisive but these recorrections actually (often) reflect the sort of changes in species definitions and technologies described above. For example, in 1923 the Common Galllinule (Gallinula galeata) of North America was lumped with the Common Moorhen (Gallinula chloropus) of Europe because they were thought, based on their breeding ranges and morphologies, to be subspecies. Then, in 2011, those species were split again, recognizing the differences in their vocalizations, bill and shield morphologies, and mtDNA.

Common Gallinule (L) and Common Moorhen (R)

If the trend shown in the graph continues—and there is no reason to expect that it will not—the number of splits will soon surpass the number of lumps. Recently, George Barrowclough and colleagues have speculated that the level of splitting resulting from the application of genomic tools and the phylogenetic species concept will almost double the number of species recognized worldwide—to 18,000—before too long. The implications of this for conservation, for understanding the evolutionary history of birds, and for my life list, are huge.


  • American Ornithologists’ Union. 1995. Fortieth supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 112: 819-830.

  • Barrowclough GF, Cracraft J, Klicka J, Zink RM (2016) How many kinds of birds are there and why does it matter? PLoS One. 2016; 11: e0166307. PMID: 27880775
  • Chesser RT, Banks RC, Barker FC, Cicero C, Dunn JL, Kratter AW, Lovette IJ, Rasmussen PC, Remsen Jr JV, Rising JD, Stotz DF, Winker K. (2011) Fifty-Second Supplement to the American Ornithologists’ Union Check-List of North American Birds. Auk 128: 600-613
  • Johnson NK (1980) Character variation and evolution of sibling species in the Empidonax difficilis-flavescens complex (Aves: Tyrannidae). University of California Publications in Zoology 112: 1-151.

  • Mayr E (1982) The Growth of Biological Thought. Cambridge, Mass: Belknap Press.

  • Stone W, Oberholser HC, Dwight J, Palmer TS, Richmond CW (1923) Eighteenth Supplement to the American Ornithologists’ Union Check-List of North American Birds. Auk 40:513–525.

  •  Vaidya G, Lepage D, Guralnik R (2018) The tempo and mode of the taxonomic correction process: How taxonomists have corrected and recorrected North American bird species over the last 127 years. PLoS ONE 13(4): e0195736. https://


  1. lumpers and splitters: respectively, taxonomists who favour placing similar populations under the umbrella of a single species versus those who seek to identify distinct populations as separate species. Ernst Mayr (1982, page 240) claimed that “As a general rule one can say that most taxonomic groups pass through a phase of rather intensive splitting when they are studied more actively, but that the splitting phase is reversed when the knowledge of the group reaches greater maturity.” I think this new paper by Vaidya et al. (2018) shows that statement to be incorrect and may reflect Mayr’s view that his Biological Species Concept was sacrosanct.
  2. while this is an excellent analysis the paper is technically detailed, a necessity given the changes in the way species have been defined and the changes in coverage of the checklist. I have tried to distill some of those details here but I am no taxonomist.
  3. quotation: from Vaidya et al (2018 page 2)

IMAGES: peeps by the author; graph redrawn by the author from Vaidya et al (2018, Figure 1); Flycatchers from; Gallinula spp. from Wikimedia Commons, in the public domain.

The First Textbook of Ornithology?

BY: Bob Montgomerie, Queen’s University | 7 April 2018

Despite their association with school courses, and the ready availability of information online, textbooks have long been—and still are—useful for working ornithologists. The OED says that a textbook is : A book used as a standard work for the study of a particular subject. By that definition, textbooks of ornithology have been with us for almost eight centuries, and are still immensely useful to student and professional ornithologists. Throughout my professional life I have maintained a small collection of hard-copy textbooks—statistics, ornithology, behavioural ecology, evolution, genetics, biochemistry—that I refer to all the time when writing scientific papers and articles, even though the majority of what I read is online or in PDFs.

The first ornithology textbook, arguably, was written by Frederick II of Hohenstaufen sometime around 1248 CE. Its title, De Arte Venandi cum Avibus, is often translated as The Art of Falconry, which is reasonable, but the title is more correctly translated as The Art of Hunting with Birds. As the title suggests, the book is largely focussed on methods for using falcons to hunt, but the first chapter is essentially a textbook of ornithology.


Frederick II had quite the resumé as he was consecutively King of Sicily, Germany, Burgundy, Italy, and Jerusalem, with a simultaneous stint as Holy Roman Emperor in the 30 years before he died at the age of 56. He has often been referred to as stupor mundi et immutator mirabilis (‘astonishment of the world and marvellous transformer’) and his accomplishments, conquests, writings, insights, and arrogance were all truly astonishing. As was his fathering of at least 19 children with four ‘wives’ and no fewer than eight mistresses. Frederick is said to have kept at least 50 falcons at court, and falconry was clearly his favourite hobby, after procreation.

From Manfred’s illuminated manuscript

Frederick’s original manuscript (a codex) disappeared during the 1248 Siege of Parma, Italy, but there are several early copies of that original—in two different versions, one of two ‘books’ and the other of six—now held in at least 12 libraries in Europe. The earliest of these were commissioned and probably edited by Frederick’s son Manfred, who was also a keen falconer.

All of those early copies are manuscripts, handwritten codices. One of the best-known is an illuminated codex of the two-book version, now in the Vatican Library. The pictures from that illuminated manuscript are often shown in books and articles about the early history of ornithology [1]. The text of the six-book version is presumably better known for it has been translated into French, German and English over a 700-year period after the original Latin codex was written.

In the six-book version, the first book is a general treatment on the anatomy and lives of birds, describing Frederick’s extensive observations and experiments. Frederick was truly a ‘Renaissance Man’, about 300 years before his time, as he based his writings on direct observation of the natural world rather than relying on an ‘authority’ like Aristotle (with whom Frederick often disagreed). When he could not make a direct observation himself, Frederick consulted with experts and often used inductive reasoning to make his conclusions.

Frederick was an inveterate experimenter and his experiments with humans were horrible, to say the least. He also tried to see if chicken eggs could hatch by the heat of the sun alone [2], and whether blindfolded vultures could find food, to see if they had a keen sense of smell. Like Audubon [3], six centuries later, Frederick concluded, incorrectly, that:

A vulture is not attracted to his carrion food by his sense of smell, although some writers maintain that he is, but relies on his eyesight. We have ourselves many times experimented and observed that an assemblage of seeled [eyelids stitched shut] vultures, whose noses were not stopped up, did not scent the meat cast before them. [4]

Frederick’s book 1 of the six-book edition is a remarkable, and remarkably accurate, treatise on the ecology, behaviour and anatomy of birds in 56 chapters. In chapter 46 (Of the Colors of Avian Plumage), for example, he notes that some species change colour with age, that some species change the colours of their plumage and soft parts for the breeding season, that herons acquire a powder down just prior to mating, and that there are ecological differences between precocial and altricial offspring.

The remainder of the book focuses on falcons, with an introduction to book 1 about falconry as a noble pursuit, book 2 on how to catch and train birds of prey, book 3 about the different kinds of lures used by falconers, book 4 on how to hunt cranes with Gyrfalcons, book 5 on hunting herons with Sacre Falcons, and the final book on hunting water birds with Peregrines.

Frederick fought wars with the Pope and was variously labelled a heretic, the antichrist, and the devil. Not surprisingly, then, his work was suppressed (hidden?) by the Catholic Church and remained virtually unknown to those interested in birds for the next 350 years. In the early 14th century, the two-book codex was translated into French, and in 1596 the Latin version was first printed and made more widely available. It was not, however, even mentioned in the ornithological works of Gesner and Aldrovandi in the 16th century, and gets only a brief quotation in the 17th in the descriptions of falcons in Ray’s Ornithology.

Title page of Wood and Fyfe 1943

Frederick’s codex was rediscovered in 1700s when one Latin edition and two German translations of the 2-book version were produced, including a copy with useful notes by Johann Gottlob Schneider in 1788 [5]. Even then, Frederick’s work was largely ignored, getting only a passing mention in Newton’s Dictionary of Birds in the late 1800s. Then, in the 1930s, Marjorie Fyfe and the indefatigable Casey Wood embarked on an English translation of the Latin six-book version. That translation and much additional material was published ina book in 1943, a year after Wood died. This is a remarkable book that deserves a separate essay of its own.

So was The Art of Falconry really the first textbook of ornithology? Looking at the content you would have to say ‘yes’. But is it really a textbook if nobody read it or used it for study (‘If a tree falls in a forest…?’). By that criterion the answer would have to be an emphatic ‘no’, and John Ray’s Ornithology would be a better candidate for the title of first ornithology textbook. The first edition of Ornithology was published in Latin in 1676, presumably in that language because it was intended for scholars. At the urging of his colleagues, however, Ray immediately produced a somewhat expanded English version in 1678, a version that became THE textbook of ornithology for the next 200 years, and is still worth reading today. Wood and Fyfe’s translation of Frederick is also worth reading and the 1961 reprint is still available on Amazon at $100 US.


  • Aldrovandi U (1599) Ornithologiae hoc est de avibus historiae. Bologna: Apud Franciscum de Franciscis Senensem.

  • Birkhead TR (2008) The wisdom of birds: an illustrated history of ornithology. London: Bloomsbury.

  • Birkhead T (2012) Bird Sense. UK: Bloomsbury.

  • Brunner B (2017) Birdmania: A Remarkable Passion for Birds. Vancouver: Greystone Books.

  • Gesner C (1555) Historia Animalium. Liber 3 qui est de avium natura. Frankfurt: Andreae Cambieriano.

  • Haskins CH (1921) The “De Arte Venandi cum Avibus” of the Emperor Frederick II. English Historical Review 36: 334 – 355.

  • Ray J (1676) Ornithologiae libri tres: in quibus aves omnes hactenus cognitae in methodum naturis suis convenientem redactae accuratè descripbuntur, descriptiones iconibus. London: John Martyn.

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

  • Walters M (2004) A concise history of ornithology. London: Christopher Helm.

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


  1. pictures from the illuminated manuscript: in Walters (2003), Birkhead (2008), Brunner (2017)
  2. eggs hatching from heat of sun: see Wood and Fyfe 1943 page 53. Haskins (1921: 342) suggests that the experiments were conducted on ostrich eggs, but Frederick makes it clear here that he was (incorrectly) certain that ostriches did not incubate their eggs for fear of breaking them, and instead they were incubated by the sun. He goes on to say that in Egypt the eggs of the barnyard fowl had been observed to be incubated by the heat of the sun. He says he summoned experts from Egypt to test this but he does not record what they discovered.
  3. Audubon’s experiments with the vulture’s sense of smell: see Birkhead (2012)
  4. quotation about vultures: from page 22 of the English translation by Wood and Fyfe (1943)
  5. Latin edition and German translations: see Wood and Fyfe (1943: lvii)