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, Paris, 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)

Red Eggs


BY: Tim Birkhead, University of Sheffield | 30 April 2018

Common guillemot eggs on Skomer, Wales showing some of the variety

The cover or frontispiece of almost every book about birds’ eggs is adorned with a picture of a common guillemot (Common Murre [1]) egg. Why? There are several reasons—the common guillemot’s egg is an extraordinary shape, it’s brightly coloured, and the variation in the colour and maculation seems almost infinite.

Egg collectors, or oölogists as they called themselves, accumulated collections of eggs mainly for aesthetic reasons, and there are few eggs more aesthetic of those of the guillemot. Collectors also liked to display the range of variation in egg colour. With the exception of one or two species like the tawny-flanked prinia Prinia subflava [2], there is more variation in the colours and patterns of guillemot eggs than any other species.

That variation is apparent if you open a museum draw of guillemot eggs. But it is a fraction of the variation apparent in freshly laid eggs on the ledges where these birds breed. Eggs in museum collections often fade unless protected from the light, and I have seen many an old collection where almost all the eggs have converged to a pale, muddy, bluish-green background colour. The markings, which are always darker, seem not to fade, so much of the variation in guillemot one sees in museums is in those maculations. Oölogists described those maculation as ‘salt and pepper’, ‘black cap’, ‘lose cap’, ‘shorthand’, and ‘scrawl’. Ornithologists have been more pragmatic and less poetic.

The majority of common guillemot eggs have a blue, green or white ground colour over which lie the various forms of maculation. In their competitive quest for eggs, oölogists sought—and sometimes fought over—the rarest types. This was especially true at Bempton Cliffs on the Flamborough Headland on England’s northeastern coast, where egging was an industry in the mid-nineteenth and mid-twentieth century. Tens of thousands of guillemot eggs were collected at Bempton each year, mainly for food, but also for the oölogists’ cabinets. Providing you were a wealthy enough oölogist, these were easy pickings. Farm labourers, known locally as ‘climmers’ (climbers) did all the work and took all the risks, scaling the 400-foot vertical cliffs to get the eggs. The collectors merely patrolled the cliff tops waiting to see what the climmers brought up.

Some Bempton ‘climmers’ in 1911 with some of the eggs they collected. William Wilkinson is in the middle.

It was a market: the climmers knew what the collectors wanted, and negotiated hard for a decent price. Farm labourers’ salaries were low and selling eggs was an important source of additional income for the climmers.  At a time when £3 was a decent weekly wage (for anyone, let alone a farm worker), an unusually coloured egg that sold for £1 was real bonus. The commoner egg shapes and colours went for a few pennies. The rarest guillemot eggs of all were those of a port-wine or blood red colour. The esteem with which such eggs were held was celebrated by the nick-name ‘Bempton Belle’ given to one egg from Bempton. Strictly speaking, the name should have referred to the female that laid this red egg since female guillemots always produce the same type and colour of egg.

When Bob Montgomerie and I visited the fabulous egg collection at the Delaware Museum of Natural History (DMNH) [3]  we found the following typewritten poem tucked into the bottom of one of the small egg boxes in a drawer full of guillemot eggs:


Listen Boys until I tell,
Of the famous egg — the Bempton Belle.
Seventeen days in June had gone;
When Jack the Climber came upon,
And brought from off the cliffs the Red;
The best ever seen by the veteran Ned.
William the Jumper struck with awe;
Said that a better egg he never saw.
Even silent Edwin Colley speaks,
Of the shape, and beautiful streaks.
The Boys all wondered at the shell,
Of the famous Red, the Bempton Belle.
Collectors who came from far and near,
Said they had never seen its peer.
Every man was fair to own,
The Bempton Belle the best egg known.
Jack the Here of the find;
Saved the egg for friendship kind.
Remember Boys the lesson taught;
True friendship can not be bought. [4]

This particular egg was part of Jeremiah Goodall’s egg collection (now in the DMNH) that he accumulated in the early 1900s. We don’t know whether it was Goodall who wrote the poem, but someone has added in pen that ‘Jack’ was Jack Hodgson, ‘Ned’ was Ned Hodgson and ‘William the Jumper’ was William Wilkinson, all climmers.

The Bempton Belle (left) now quite faded, and a 3D-printed replica (right) painted to represent how the Bempton Belle might have looked when it was fresh [5].
The colour of all birds’ eggs is determined by just two pigments, biliverdin and protoporphyrin IX. Our analyses show that red guillemot eggs contain only the latter pigment, as one might expect. Some other birds, and especially corvids—which typically lay greenish or blueish eggs—occasionally lay entire clutches of red eggs, referred to as ‘erythristic’, and these too were popular with egg collectors. Red guillemot eggs are essentially erythristic, albeit with heavy maculation.

We have attempted to assess just how rare such eggs were, and after looking at our own data and those of other guillemot researchers, we estimate that fewer than one in 1000, or possibly 10,000 female guillemots produce such extraordinary eggs [6].


  • Birkhead TR, Montgomerie R (2018) Rare red eggs of the Common Guillemot: birds, biology and people at Bempton, Yorkshire, in the early 1900s. Archives of Natural History 45: 69-79.
  • Boehm EF (1950) Abnormal erythrism in birds’ eggs. Emu 50:139.
  • Caves EM, Stevens M, Iversen E, Spottiswoode CN (2015) Hosts of brood parasites have evolved egg signatures with elevated information content. Proceedings of the Royal Society of London B 282: 20150598.
  • Gill F, Donsker D, Eds (2018) IOC World Bird List (v8.1). doi: 14344/IOC.ML.8.1. Online at
  • Gross AO (1966) Erythristic eggs. The Wilson Bulletin 78: 127–128.
  • Rarer ECS (1918) IV.‐Erythrism in bids’ eggs: an address read at the Third Oological Dinner on 26 September, 1917. Ibis 60: 68–75.


  1. common guillemot: The ‘official’ name of this species (Uria aalge) is Common Murre according to Gill and Donsker (2018) and HBW Alive/Bird Life International. Common guillemot is the name still used for this species in the UK.
  2. prinia egg colours: see Cave et al. (2015)
  3. Delaware Museum of Natural History: just north of Wilmington, this museum was established
  4. poem: See Birkhead and Montgomerie (2018) for a picture of the original. On that typescript someone has added some details in pen, and changed the last line and added two more: Honest Jack could not be bought/To your children the story tell/Of Jack the Climber, & the Bempton Belle.
  5. Bempton Belle photo: the actual egg now in the collection at DMNH; I painted the replica to show what the original probably have looked like, based on red guillemot eggs currently in the collection of the Natural History Museum at Tring (see Birkhead and Montgomerie 2018, Figure 1)
  6. rarity of erythristic guillemot eggs: see Birkhead and Montgomerie (2018) on the Bempton Belle and erythrism in murre eggs. See also Boehm (1950), Gross (1966), and Rarer (1918) on erythrism in birds’ eggs in general. Erythrism is either the result of addition protoporphyrin IX being added to the shell by the female, of biliverdin not being added to a shell that normally would have both pigments.

IMAGES: Bempton ‘climmers from an old postcard; Bempton Belle poem and photo © Delaware Museum of Natural History, used with permission of Dr Jean Woods, Curator of Birds; photo of guillemot eggs on Skomer and replica red egg © T R Birkhead.

Serendipity 101

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

The more I read about the history of ornithology, the more it strikes me how important serendipity—blind luck—has been to that history. Ernst Mayr’s career, for example, was a long series of fortunate events. No question that Mayr was brilliant, ambitious and creative, but the goddess of fortune was definitely smiling on him.

One hundred and twelve years ago today, San Francisco lay in ruins, decimated [1] by 4 days of fires that followed the earthquake that struck in the early morning on 18 April 1906. The earthquake itself was monstrous but it was the fires—fuelled by broken gas mains and the largely timber construction of much of the city—that wreaked the most havoc for four consecutive days following the initial quake. Charles Richter was only 6 years old in 1906—and thus had not yet invented his eponymous scale—but geologists figure that the Great 1906 San Francisco Earthquake  would have numbered 7.9 on that scale.

The fires of April 2006, San Francisco
Guadalupe Storm Petrel, painting by Keulmanns 1906

The California Academy of Sciences was destroyed by the quake and most of its specimen collections burned in those fires. Nonetheless, a particularly valuable portion of the herbarium [2] was rescued by the heroics of Alice Eastwood, the curator of botany. The museum director Leverett Loomis was able to rescue only two bird books and two bird specimens (both of the Guadalupe Storm Petrel [3]) but all of the other ornithological material was lost.

While the Cal Academy’s bird collection was lost, there was a serendipitous silver lining in that cloud of smoke. In 1904, Loomis had commissioned Rollo Beck to lead a 17-month-long collecting expedition to the Galápagos. Beck was a superb collector who lived in California making extensive collections of birds along the west coast of North America for museums and private collectors like Sir Walter

Galápagos expedition members 1905 (Beck is seated in the middle)

Rothschild, who figures prominently in Ernst Mayr’s story of success. Loomis, however, had a ton of trouble trying to find a suitable ship to either buy or lease for that expedition. So the departure date had to keep being advanced from October 1904 until the end of June 1905 when they finally were able to leave San Francisco on the rebuilt schooner Academy. If the expedition had left in 1904, as originally planned, it would have returned in March 1906 just in time to be burned by the fires of April.

Thus, in April 1906, when the Cal Academy collections burned, Beck was still in the Galápagos. He returned in November 1906 bringing with him more than 78,000 specimens—including 8688 birds (3200 of which were Galapagos Finches [4]) and 2000 birds’ eggs —that would form the nucleus of the collections in the rebuilt museum [5]. The Cal Academy restored the museum and reopened in 1916, then rebuilt the whole structure with improved earthquake and fire protection in 2008. It is a magnificent museum, aquarium and small zoo today and well worth a visit next time you are in San Francisco. It also still holds the largest collection of Darwin’s Finches in the world. Those specimens formed the basis for much of the work that Robert Bowman and David Lack did on the anatomical adaptations of those birds [6].

California Academy of Sciences, Golden Gate Park, San Francisco 2008

Beck was later hired by the American Museum of Natural History to head the Whitney South Sea Expedition to the south Pacific islands to collect birds and anthropological material. Beck set out with his crew in 1923 but in 1929 he got very sick and had to go home. That left the expedition without an ornithologist. Sir Walter Rothschild knew

Ernst Mayr (right) in New Guinea 1928

about the 24-year-old Ernst Mayr then working as an assistant to the great German ornithologist Erwin Stresemann in Berlin. Rothschild had promised Mayr a collecting trip to the tropics on completion of his PhD in 1922 but no opportunities had presented themselves until 1928 when he was able to send Mayr to New Guinea. Thus Mayr was in the right place at the right time when the Whitney expedition needed a new ornithologist so Rothschild asked him if he could take over to complete the work that Beck had started. After returning to Europe in 1930, Mayr moved to New York to work on the specimens collected on the Whitney expedition and established his early career with those publications. And the rest, from those very lucky beginnings, is history.

The take home lesson from Serendipity 101 is that ‘shit happens’, but good things happen too and there is really nothing we can do about many of those devastating events like earthquakes. What we can do, as Ernst Mayr’s life so amply demonstrates, is to recognize and be prepared to take advantage of the good things, and to try not to be too discouraged when the goddess of fortune [7] takes a holiday.


  • Bowman RI (1961) Morphological differentiation and adaptation in the Galapagos finches. University of California Publications in Zoology 58:1–302.
  • Godman FDC (1907) Monograph of the Petrels (order Tubinares). London: Witherby & Co.

  • James MJ (2012) The Boat, the Bay, and the Museum: Significance of the 1905-1906 Galápagos expedition of the California Academy of Sciences. Pages 87-99 in Wolff M, Mark Gardener M (editors) The Role of Science for Conservation. London: Routledge
  • Lack D (1945) The Galápagos finches (Geospizinae): A study in variation. Occasional Papers of the California Academy of Sciences 21:1–159.
  • Lack D (1947) Darwin’s Finches. Cambridge: Cambridge University Press.
  • Lowe P (1936) The finches of the Galápagos in relation to their evolution. Ibis 1936:310–321.


1. Decimated: literally reduced to 1/10th of its former size as 90% of the homes were destroyed. See here  for a summary of the effects of those fires

2. Valuable portion of the collection: these were type specimens that Eastwood had kept separate from the main collection. This was an unusual practice in those days but proved to be lucky as she was able to grab them in a few minutes. On arriving at the museum, she found that the marble staircase had collapsed so she climbed to the 6th floor herbarium on the iron railing. She then gathered up those type specimens with her friend Robert Porter. Eastwood then climbed back down the bannister to the ground floor to gather up the specimens that Porter lowered out the window, on a rope that they had put together from scraps. See here  for more details.

3. Guadalupe Storm Petrel: although this bird was still considered to be abundant on its Guadalupe Island (Mexico) breeding grounds in 1906, it was being heavily preyed upon by cats that were introduced to the island in the late 1800s. The last two specimens were collected in 1911 and the last breeding was recorded in 1912. The species was never seen again (see here for more details). Loomis must have taken these two because they were the type specimens as he could not have guessed that the species was soon to be extinct. The painting by JG Keulmans is from Godman (1907)

4. Galápagos Finches: were not normally called Darwin’s Finches until Percy Lowe (1936) used that term

5. Galápagos expedition of 1905-06: see James (2012) for details

6. Lack and Bowman studies of Darwin’s Finches: see Lack (1945, 1947) and Bowman (1961)

7. Goddess of Fortune: Tyche to the Greeks, Fortuna to the Romans

IMAGES: all from Wikipedia, in the public domain

A Bird’s Eye View

BY: Bob Montgomerie, Queen’s University | 17 April 2018

When I started my PhD at McGill University, in 1973, I was thrilled to discover that the university had a library devoted just to ornithology: the Blacker-Wood Library of Zoology and Ornithology. This library was a primarily large room located in the basement of the university’s  immense Redpath Library just down the road from the Biology Building. Besides a fabulous collection of contemporary books and journals about birds, the library also had a rare book room full of treasures like the feather book of Dionisio Minaggio. It also had a hand-coloured copy of John Ray’s Ornithology of Francis Willughby published in 1776 and ostensibly given as a present to Samuel Pepys, the famous 17th century diarist [1]. 

I remember the librarian, Eleanor MacLean, telling me that the library subscribed to more than 100 ornithological journals and newsletters. That seems like a lot but they did get the separate bird journals published by many of the provinces in Sweden, as well as a dozen or more in foreign languages that I could not read, so that total might not have been far off. It was a real treat for me to spend every Friday morning poring over the latest acquisitions. While I love the convenience of the internet, and rapid search engines, there was something oddly rewarding about opening the latest issues of journals from around the world, all devoted to the study of birds.

That library was founded in 1920 by Casey Wood, a Canadian-American [2] ophthalmologist who was also an avid—OK, obsessive—book collector. On Friday (20 April), I will be going to Montreal to participate in a symposium celebrating his contributions to the library in particular and to ornithology in general.

Wood wrote several papers about birds but his signal ornithological work was a little-known volume called The Fundus Oculi of Birds, published in 1917. Because Wood was an ophthalmologist and immensely curious, he thought it might be interesting to use the tools of his trade to study the eyes of birds. To do that, he employed the techniques that he had used in his practice, and further developed some procedures for studying the eyes of birds. His main interest was the structures of the fundus oculi, at the back of the eye—the retina, the fovia(s), the nerves and the pectin.

2018-04-16 21.11.16
Wood adapted the methods he used to strudel the eyes of his patients (left) to examine the fundus oculi of birds (right). From Wood (1917)

64BCC83D-A67F-46C9-8914-7F692BDF0D6AWood realized early on that his observations would be most useful if he could somehow illustrate what he was seeing through his ophthalmoloscope. In those days, photography was simply not advanced enough to provide enough depth of focus:

In spite of recent advances in that direction, attempts to reproduce the colored (ophthalmoscopic) appearances of the fundus by photography have so far failed. [3]

Instead he employed the talents of Arthur William Head [4], a celebrated UK artist, to draw and paint the fundus oculi of more than 80 bird species. Wood does not record how Head did this—did Wood train Heath to use the opthalmoscope or did he roughly sketch what he saw and then work with Head to make Head’s illustrations match what Wood saw? Wood says that it takes a lot of experience to be able to use the ophthalmoloscope but maybe he was able to teach Heath his techniques. Wood also says that his methods allowed only a short period of observation of the fundus oculi so it is remarkable how detailed these illustrations are, however they were produced.

file 2018-04-16, 21 25 12.png
Arthur Head’s paintings of the fundus oculi of Great Black-backed Gull (left) and African Penguin (right) showing the pectin (black) and retina (red or brown). From Wood (1917)

Perhaps surprisingly, Wood took a very modern approach to his subject, as he attempted to examine at least one species in each avian order to see if there were patterns related to the bird’s way of life—an early example of what we now call comparative analysis.

 The oculi of Birds exhibits a great variety of areas of distinct vision, and these correspond closely to the habits and habitat of these animals especially their methods of obtaining food, of escape from enemies, of migration, of reproduction, etc. [3]

But he also wanted to determine whether the fundus oculi might be a useful taxonomic tool, helping to define the evolutionary relationships among species. To that end, he sampled and illustrated the fundus oculi of birds from 24 of the Orders of birds that were recognized in those days, missing only 5 of the Orders that we (or at least some authorities) recognize today (Turniciformes, Craciformes, Gabuliformes, Coliiformes and Musophagiformes). There was a bit of a taxonomic signal there, in Wood’s opinion, but not clear enough to be useful. 

Side view of the eyes of an Ostrich (left) and  Kiwi (right) showing a clear difference in pectin (black) structure that Wood thought might be related to visual acuity, with the kiwi needing less as it hunts mainly by smell at night. From Wood (1917)

 Instead, Wood showed nicely that the pectin size and shapevwere likely related to visual acuity, and that some birds have two fovias, related either to visual acuity or more likely the need to switch from binocular to monocular vision while foraging. Even today, a century later, we still do not have definitive studies of these traits in the fundus oculi, as far as I can tell.

 Wood also found that the pectin degenerated in domesticated birds suggesting to me that it is a costly structure that becomes smaller under relaxed selection when visual acuity is no longer important for foraging or survival.

 Reading Wood’s Fundus Oculi Book for my talk in Montreal reminded me once again of the tremendous value in reading the older literature once in a while. I have been interested in avian vision for about 25 years but had never really thought or read about the pectin before. To the best of my knowledge, many of the patterns that Wood described have not been further investigated even though we now have all of the tools needed to measure and quantify the structures he described and illustrated. Wood’s book is not an easy read as it is largely descriptive and lacks focus (no pun intended). There are enough gems buried there, though, that anyone interested in the vision of birds might want to have a look at this interesting book.


  • 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  DOI 10.1007/s10336-009-0413-3
  • Wood CA (1917) The Fundus OCuli of BIrds ESefially as Viewed by the Ophthalmoloscope: A Study in Comparative Anatomy and Physiology. Chicago: Lakeside Press. Available here


1. Pepys’s  copy of Willughby and Ray’s Ornithology: see Montgomerie and Birkhead  (2009)

2. Casey Albert Wood: (1856-1942) was born in Wellington, Ontario, attended high school in Ottawa, and got his medical degrees from McGill University. He eventually settled in Chicago where he practiced ophthalmology. See here for details

3. Quotations: from Wood 1917 page 7

4. Arthur William Head: (1861-1930) was a celebrated artist who lived in London, England, see here for examples of his work

The Utmost Harmony

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

I gave my first research talk at a ‘big’ international conference at the AOU meeting at Haverford College (Pennsylvania) in 1976. I talked about my work on Mexican hummingbirds and I was nervous, in part because Frank Gill—who was then doing great work on hummingbirds—was talking right before me. The chair of our session was some old guy with unruly grey hair. Though I really wanted to hear Frank’s talk, I was too busy thinking about what I was going to say to actually pay attention to his words. Until, that is, when he finished answering a couple of questions. As he was about to leave the stage, Frank said to the chair “Thanks, Ernst”. “OMG”, I thought [1], “that’s Ernst Mayr, and I may not survive this.”

Ernst Mayr ca 1976

I completed my presentation on autopilot, out of sheer terror. But Professor Mayr asked me a couple of excellent questions and thereby, very graciously, put me at ease and made my presentation seem like a success. I had been to a few AOU meetings before but that incident convinced me that I had found my academic home. It seemed that everyone from the most famous—Mayr and Gill—to the greenest student (me) could talk about birds in an environment characterized by the utmost harmony.

Indeed, that’s how someone [2] described the first AOU meeting, in a report in the Bulletin of the Nuttall Ornithological Club in 1883:

The session of the Convention occupied three days, and was marked throughout with the utmost harmony; at adjournment…hearty expressions of satisfaction with the results of the session were heard from all who had shared in its deliberations. The general good feeling rose to a degree of enthusiasm auguring well for the future work and prosperity of the Union, the organization of which, under such auspicious circumstances, cannot fail to mark an important era in the progress of ornithology in America. [3]

He was right, that first convention did augur well for the future work of the ‘Union’—and it has gone from strength to strength over the past 135 years. The AOU was the gifted child of the Nuttall Ornithological Club in 1883 and the proud parent (along with the Cooper Ornithological Society) of the American Ornithological Society (AOS) in 2016.

El Conquistador Hotel, Tucson, Where the 2nd AOS meeting is about to begin on 11 April 2018

Today I am in Tucson waiting for the start of the 2nd annual AOS meeting. I am anticipating some great science, some reconnecting with old friends, and seeing some interesting birds. I am not really a birder and especially not a twitcher, so I will not make the trek to Madera Canyon to see the Elegant Trogon reported there last week. This morning though, I watched  a pair of Verdins building a nest, and spent an hour in the midst of a dozen pairs of courting Great-tailed Grackles. I always prefer watching behaviour over searching for rarities. But I digress.

That first AOU meeting in 1883 was the result of a letter sent on 1 August 1883 by three officials [4] of the Nuttall Ornithological Club to 46 American and 2 Canadian ornithologists,


You are cordially invited to attend a Convention of American Ornithologists, tiobe held in New York City, beginning on Seotember 26, 1883, for the ourpose of founding an AMERICAN ORNITHOLOGISTS UNION, upon a basis similar to that of the “British Ornithologists Union”…The object of the Union will be the promotion of social and scientific intercourse between American ornithologsts, and their cooperation in whatever may tend to the advancement of Ornithology in North America…” [3]

Twenty-one men attended the 3-day conference in the library if the American Museum of Natural History. They declared themselves to be founders of the AOU, along with Spencer Fullerton Baird and J A Allen who were unable to attend. Their first order of business was to establish a constitiution for the new society followed by the election of other ornigthologists—all men as far as I can tell—to various classes of membership: 21 Foreign, 20 Corresponding, and 81 Associate in addition to the 47 Active members that included the Founders.

They also established six committees that nicely reflected 5 of the major ornithological interests of the day: Classification of North American Birds, Migration, Avian Anatomy, Oölogy, and Faunal Areas. The sixth committee was charged “to investigate the eligibilibity or ineligibility of the European House Sparrow in North America”. I do not know what that sixth committee eventually decided, but it is clear from events over the next century that the House Sparrow did not care.


  • Anonymous (1883) The American Ornithologists’ Union. Bulletin of the Nuttall Ornithological Club 8: 221-226


1. OMG: I did not, of course, think that in 1976, as it did not enter the slang lexicon for another 20 years

2.  someone described that first AOU meeting: see Anonymous (1883); normally i would have assumed it was the editor of the journal but that was J A Allen who was ill and could not attend the meeting

3. quotations from Anonymous (1883)

4, three Nuttall Club officials: were J A Allen (editor of the Nuttall Bulletin), Elliott Coues (associate editor) and William Brewster (President of the Nuttall Club)

¿Hay huevos?

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

I did my PhD field work in Nayarit, Mexico, mainly in the coastal town of San Blas. On our first long drive there from Montreal, my fellow PhD student—Neil Brown—and I attempted to learn some basic Spanish, using a small Berlitz guide. Neil dropped me off just outside town, then headed off to his own study area near Tepic, in the mountains. I soon found a place to stay and went to the local tienda to get food. Wanting to try out my new Spanish, I asked the young male owner “¿Tiene huevos?” to which he gave a sheepish smirk and pointed to a basket on the counter full of fresh eggs. After a couple of weeks of this, he said he’d help me learn some Spanish, beginning with the proper way to ask for eggs (¿Hay huevos?)—I was really asking him if he had any testicles. My Spanish has not much improved in the intervening 45 years but my scientific interest in both eggs and testes has vastly expanded.

ONRS nest record card

Like generations of teenage boys before me, I collected birds’ eggs. Unlike most of them, however, my collection was almost entirely photographic. In 1965, while still in high school, I had the good fortune to meet George Peck, a veterinarian who had just volunteered at the Royal Ontario Museum to coordinate the Ontario Nest Records Scheme (ONRS). This ‘scheme’ began in 1956 as a collection of file cards, one per nest, summarizing details about the location, clutch or brood size, and, sometimes, the history and fate of a nest found in Ontario. The ONRS was modeled after a similar scheme run by the British Trust for Ornithology.

George Peck was then an avid birder and (former) egg collector who had made it his lifelong hobby to photograph the nest and eggs of every North American bird [1]. For several years we spent many a weekend searching for nests throughout southern Ontario, photographing their contents and, often, setting up blinds to watch and photograph the parents as well. By the mid-1960s egg-collecting was illegal in Ontario [2] so we only took (and preserved) eggs from abandoned nests. George was always extremely careful around birds’ nests to minimize the risk of disturbing the parents and he taught me a lot about being a careful observer and note-taker.

The exquisite nest and four eggs (plus one cowbird egg) of a Blue-gray Gnatcatcher, high in a maple tree at Rondeau Provincial Park, Ontario, in 1969 (R Montgomerie Photo)

Today the ONRS contains more than 120,000 nest records and provides a superb historical record of the distribution and other details (that George liked to call ‘nidiology’) of breeding birds in Ontario. Peck and James used these records to compile the first distributional and nidiological survey of breeding birds in Ontario, an early and quite successful product of citizen science.

I am writing this post on Easter Egg Weekend, at a time when we are seeing something of a renaissance in the scientific study of birds’ eggs. Just last Saturday (31 March), BBC TWO aired a special on The Wonder of Eggs, hosted (of course) by David Attenborough [3]. And last year, birds’ eggs were on the cover of Science, with a comprehensive paper on the evolution egg shape by Cassie Stoddard (Princeton Univ) and her colleagues. Recent books by Tim Birkhead (Univ Sheffield) in 2016 and Mark Hauber (Univ Illinois) in 2014 have respectively highlighted both the exquisite biology and the amazing diversity of bird’s eggs.

As a budding oölogist [4], I devoured the Romanoff’s 1949 classic The Avian Egg, but by the time I started graduate school in the 1970s research on birds’ eggs had largely fallen by the wayside. I suspect that the scientific study of wild birds’ eggs was a victim of the demonization of egg-collecting by conservation groups and the very strict laws that eventually became established in Europe and North America. In some places, these laws even make the private curation of an egg collection illegal and have lead to the—in my opinion senseless—destruction of countless egg collections.

huicholFGDLCFreud would undoubtedly have had a field day speculating on the association between egg-collecting and adolescent males. But I suspect he would have been wrong, as he was in much of what he wrote. While I was doing my PhD field work in San Blas, I met a local Huichol artist named Fermin Gonzalez de la Cruz who sold me a couple of his excellent yarn paintings. The details of one are shown to the right and were featured on the cover of National Geographic several years ago.

Huichol_shamanMany of the Huichol men that I encountered wore sombreros adorned around the rim with trinkets, and one of those men’s trinkets were mummified hummingbirds. I was studying hummingbirds, so this really intrigued me. When I asked him about it, he told me that hummingbirds were the traditional sombrero ornaments in his culture [5]. They were hard get, he said, so they became a sign that the bearer was a good hunter. As a budding behavioural ecologist, I interpreted that as a signal of status either to other males, or to potential mates. 

Birds eggs are one of several jewels of the bird world, not only beautiful but also delicate, hard to find, and (potentially) wearable. I do not know if there are any cultures that used bird eggs as ornaments but I do wonder if the European and North American obsession with egg collecting in the late 1800s was as much a signal of status as a desire to learn more about avian biology.

Regardless of the underlying psychology, the great egg collections now housed and curated in the museums of the world are a treasure trove of scientific and historical material—a subject that we will explore often on this site. Many avid (and, of course, male) collectors donated their large collections to—or indeed established—museums.

In Britain, the magnificent collection of more than a million eggs at the British Museum (Natural History), got a head start from an egg collector when Walter Rothschild donated his extensive collection (and his estate at Tring) in the 1930s. Similarly the superb egg collections at the Western Foundation of Vertebrate Zoology in Camarillo, California (by Ed Harrison) and the Delaware Museum of Natural History (by John du Pont) were both established by avid, even obsessive, egg collectors. In Britain, once the Protection of Birds Act [6] was passed in 1954, many private egg collections were destroyed by fearful owners, or zealous enforcers of the law [7].

Common Murre eggs at the Delaware Museum of Natural History

It is often said that we should study history to provide lessons for today. From the history of oölogy, we should have learned that egg collections: (i) are a valuable scientific resource, (ii) can help to solve some conservation problems [8], and (iii) are relatively easy to preserve. There is also no concrete evidence that the collecting of even thousands of eggs had any real impact on all but a very few very rare bird species. While I would never advocate the collecting of eggs as a hobby, as it once was, I feel that some of the current restrictions on scientific egg collecting are misguided.

When Paul Sweet of the American Museum of Natural History was asked recently how many eggs were in their collection [9], he answered, apparently without hesitation, 17,921. He knew that number precisely, he said, because it had not changed in years. That’s really a shame as there is much useful information being lost when museums do not maintain an active egg-collecting program, one that minimizes any possible damage to bird populations while maximizing the usefulness of the specimens through careful preservation, data collection and curation.


  • Birkhead T (2016) The Most Perfect Thing: Inside (and Outside) a Bird’s Egg. London: Bloomsbury.

  • HauberCoverHauber ME (2014) The Book of Eggs: A Guide to the Eggs of Six Hundred of the World’s Bird Species. Chicago: University of Chicago Press.

  • Peck GK, James RD (1987) Breeding birds of Ontario: nidiology and distribution (2). Toronto: Alger Press.

  • Ratcliffe  DA (1967). Decrease in eggshell weight in certain birds of prey. Nature 215: 208-210.
  • Romanoff AL, Romanoff AJ (1949) The Avian Egg. New York: John Wiley and Sons
  • Stoddard MC, Yong EH, Akkaynak D, Sheard C, Tobias JA, Mahadevan L (2017) Avian egg shape: Form, function, and evolution. Science 356: 1249-1254.


  1. George Peck: managed the ONRS for the rest of the century, elevating it to probably the best such set of nest records in North America. He continued to photograph nests and eggs until his eyesight failed him, coming very close to his goal of photographing all North American species. His son, Mark, now works at the ROM and continues his father’s interest in eggs and photography.
  2. egg collecting illegal in Ontario: as far as I can tell, egg collecting actually became illegal across North America, at least for migratory birds, with the Migratory Birds Convention Act of 1918. Apparently, that part of the act was rarely, if ever enforced with respect to the hobby, until the 1050s at least
  3. Attenborough’s Wonder of Eggs: has not yet aired in North America but you might be able to watch all of it (if you are in the UK) or some short clips on the BBC TWO website, and you can read about it here
  4. oölogist: literally means ‘one who studies’ eggs, but the term has been applied only to birds’ eggs; during the late 1800s and early 1900s ‘oölogist’ meant ‘egg collector’ so the term is not used very often today, except by scientists
  5. hummingbirds on sombreros: Some recent sources suggest that this story is apocryphal (though my story is true); while the hummingbird was imbued with mythological powers by the Huicholes, they apparently adorned their sombreros with a wide variety of natural objects in prehistoric times but now, like satin bowerbirds, use readily available man-made objects as decorations
  6. Protection of Birds Act of 1954: you can read a little bit about it here; I suspect that the passing of this Act had some influence on egg-collecting in North America, as well, even though egg-collecting had already been ‘illegal’ here for 36 years.
  7. destroying egg collections: Freud would undoubtedly have had a field day speculating on the psychology of the egg crushers
  8. egg collections solving conservation issue: the measurement of the eggs of some species in collections was key in pinpointing the effects of DDT on declining Peregrine Falcon populations (see Ratcliffe 1967)
  9. number of eggs in AMNH  collection: quoted in an article in The Atlantic

IMAGES: Huichol shaman from Wikipedia; all other photographs by the author

Ornithology, March 1918

BY: Bob Montgomerie, Queen’s University | 26 March 2018

Unpublished manuscript dated March 26th, 1918
by Lloyd Kerswill, King City, Ontario [1]

It’s nearly the end of March and I have in front of me the first issues for 1918 of the world’s major ornithological journals—The Ibis, The Auk, The Condor, The Wilson Bulletin, The Emu, and Journal für Ornithologie. Even though there are probably only a few thousand professional and amateur ornithologists worldwide, the number of scientific publications about birds has burgeoned this century. Several journals do a nice job of publishing information on some current papers, but I thought it might be useful to provide a general overview here.

Even just summarizing all of the books—let alone the scientific papers—about birds in any of the great libraries is a daunting task that may soon be impossible.  I know that Messrs. William Mullens and Kirke Swann are putting together a comprehensive bibliography of British ornithology, but even that may take years to complete [2]. The AOU has almost completed a Bibliography of Bibliographies summarizing approximately 200 bibliographies including 26,000 titles and that might see publication in the next year or two [3]. The sort of listing of all ornithological publications attempted by earlier compilers, however, is probably a thing of the past.


These are trying times for humanity. The German juggernaut has just this month crossed the Somme in this endless war, and may soon occupy all of Europe. For months, now, the front page of the New York Times has been dominated by war news. I see also that there has been an outbreak of Spanish influenza in Kansas. This deadly disease is the scourge of modern medicine as it seems to appear from nowhere then sometimes spread like wildfire [4]. The AOU is taking up a collection to support our members who have gone overseas to aid in the war effort and I encourage you all to contribute to this worthy cause. The AOU will also not require that dues be paid by members in active military service [5].

As I write, the very popular Darktown Strutters’ Ball is playing on my gramophone. This catchy song was written by fellow Canadian, Shelton Brooks, who was born down in Amherstburg near Point Pelee where we go every year to watch the spectacular spring migrations. Those new musical rhythms are uplifting in this time of world tragedy, as are the ornithological journals. There’s a lot of interesting natural history being published and we should be grateful that most scientific research transcends the boundaries of nations and politics.

Here’s a brief summary of what’s in those first issues of ornithological journals in 1918 [6], in alphabetical order:

AUKjan1918THE AUK (Jan 1918): papers and notes on species’ distributions (21), breeding biology (2), migration (1), taxonomy (2), history (1), anatomy (1), behaviour (1) plus a memorial, and notes on society business, ornithological journals, papers in other (i.e., not major ornithological) journals, publications (mainly books) received, and 18 brief notes on bird papers from the recent literature.

THE CONDOR (Jan-Feb 1918):  papers and notes on species’ distributions (12), migration (2), anatomy (1), and behaviour (1) plus 2 brief reviews of recent books, and a note from the editor (Joseph Grinnell) on some recent papers that interested him and society news including the recent deaths of members. I like the personal touch that Grinnell adds to this journal and his insights into all things avian.

EMUjan1918THE EMU (Jan 1918): papers and notes on species’ distributions (2), breeding biology (3, incl 1 on oölogy), and taxonomy (1), plus a memorial, some 3 letters from members on a variety of ornithological topics, and the annual report of the RAOU for 1917. That report includes details of the recent donation of H. L. White’s magnificent egg collection to the RAOU, as well as many other donations of egg and skin collections to the society. The RAOU will deposit all of these specimens in the National Museum in Sydney where they will be preserved for study and the education of future generations.

IBISjan1918THE IBIS (Jan 1918):  papers and notes on species’ distributions (1), breeding biology (3, and 2 of these on oölogy), taxonomy (1), history (1), anatomy (1), behaviour (1) plus a memorial, some notes on recent ornithological publications (including books and journals), and 7 letters from members on a variety of ornithological topics, as well as some ornithological news. One of those news items summarized the eggs presented at the Third Oölogical Dinner held in London last September where the main topic of discussion (and display) was erythristic eggs. The paper that I have categorized as ‘history’ lists all of the coloured plates published in The Ibis from 1859 to the present—a wonderful resource.

JOURNAL FÜR ORNITHOLOGIE (Jan 1918): papers on species’ distributions (4), and breeding biology (1), plus summaries of recent meetings of the Deutsche Ornithologen-Gesellschaft. The paper that I have categorized as ‘breeding biology’ is an interesting one on the effects of the war on birds.

THE WILSON BULLETIN (March 1918): papers and notes on species’ distributions (6), and breeding biology (1), plus a complete listing of the 54 women and 205 men who are currently members of the Wilson Ornithological Club, including my friend James H. Fleming of Toronto and 8 other Canadians.

From today’s perspective (March 2018)

parrotsI was surprised to find so little taxonomy in these journals as that was probably the main scientific pursuit of professional ornithologists in those days. Papers and notes on the birds of different places and regions dominated all of the journals. The Auk had the most papers and the widest diversity of topics but I found the papers in The Ibis and The Emu most relevant to my current interests in behavioural ecology. I was also surprised at both the number and quality of photographs in those journals.

I particularly liked a paper by Gregory M. Mathews on the plumage colours of Australian Rosellas published in The Ibis, with two beautiful plates painted by the superb Danish artist Henrik Grønvold. That paper speculated on the evolution of the different colour patterns within and between species and is full of interesting insights.

Besides the science noted above, one striking thing about The Auk in 1918 is that the editor required manuscripts to be submitted to him at least 6 weeks before the publication date. That means 6 weeks from submission to print! Presumably the editor did all of the reviewing (if any) but even that is a remarkable turnaround by today’s standards.


  • Mathews GM (1918) The Platycercine parrots of Australia: a study in colour-change. Ibis Tenth Series Vol 6: 115-127

  • Mullens WH, Swann HK (1919) A bibliography of British ornithology from the earliest times to the end of 1912, including biographical accounts of the principal writers and bibliographies of their published works. London: Macmillan and Co., Limited.


1. Article from 1918: This post intended to be my (first and rather feeble) attempt at historical fiction. My idea is to try to recreate the language and feel of something written in a century ago, based on the facts available. The fictitious writer is my real maternal grandfather, Lloyd Kerswill, who did live in on the family farm in King City in 1918 (when he was 34) but he was not particularly interested in birds. He did fuel my interest in behaviour and natural history, though, and I can remember enough about him to write in his voice and to relate to some of his lifestyle on the farm, which still exists on the outskirts of Toronto. I used to listen to his ancient one-sided, shellac-based records on his wind-up gramophone in the 1950s.

2. Mullens and Swann: This work was initially published in parts, and part 6 had just been issued by Jan 1918. That turned out to be the final part so the book was put together and published officially in 1919, and a further supplement was published as a single volume in 1923.

3. Bibliography of Bibliographies: In a brief search, I can find no further mention of this work and assume that it was never published

4. Influenza: The 1918 flu pandemic eventually killed a staggering 50–100 million people worldwide

5. No AOU dues for military: see Palmer 1981 Auk 35: 70

6. Journals from 1918: back issues of The Auk, The Condor and the Wilson Bulletin are all freely available at SORA; back issues of IbisEmu and Journal für Ornithologie are available at the BioDiversity Heritage Library

The Peculiar Etymologies of Ross’s Birds

BY: Bob Montgomerie, Queen’s University | 19 March 2018

When I was first starting to learn about birds, I was particularly intrigued (and delighted) with the peculiar names that had come from old English (dunlin, cormorant) and other languages (guillemot, eider), and those that were onomatopoeic (cuckoo, chickadeee). I also assumed that those birds named after people (Audubon’s Warbler, Temminck’s Stint, Townsend’s Warbler) were a nice touch, maybe honouring those early naturalists who had either discovered those birds or studied them in some detail.  It seems that bird names must be generally interesting to birders and ornithologists as there are more than a dozen books and papers on the origins of English (common) bird names dating back into the 1800s [1].

Ross’s Geese (painting by Angus Shortt 1948)

Recently, while putting together some material on the history of Arctic ornithology, I discovered, somewhat to my dismay, that the naming of birds after people was not nearly as rational or commemorative as I had once thought. I had assumed, for example, that the lovely Ross’s Goose had been named after Admiral Sir John Ross, the Scotsman who explored the Canadian Arctic in the early 1800s in search of a Northwest Passage with William Parry. Or maybe even after Captain Sir James Clark Ross, John’s nephew, who also explored the Canadian Arctic, but is more famous for his Antarctic exploits. Indeed, James Ross was maybe the more likely candidate as he was something of a naturalist where John Ross was not. Many famous early explorers were also naturalists who contributed immensely to our early knowledge of Arctic birds, so it seemed to me that it was reasonable to have immortalized them in the common (and scientific) names of birds.

The Ross’s Goose was actually named, in 1861, after Bernard Rogan Ross, a Hudson’s Bay Company clerk and chief trader at various settlements (forts) owned by the company in what was called, in those pre-Canada days, The North-Western Territory and Rupert’s Land [2]. Ross was a keen naturalist who sent hundreds of specimens to the Smithsonian in Washington and the British Museum (Natural History) in London, along with excellent notes on the habitats and nesting habits of each species that he collected. He also published on mammals, birds, and ethnographic topics in 1861 and 1862 [3].

The Ross’s Goose was actually ‘discovered’ almost a century earlier, in the 1760s, and given the English name ‘Horned Wavey’ in 1795, by the great Arctic explorer and naturalist Samuel Hearne, who wrote:

HORNED WAVEY. This delicate and diminutive species of the Goose is not much larger than the Mallard Duck. Its plumage is delicately white, except the quill-feathers, which are black. The bill is not more than an inch long, and at the base is studded round with little knobs about the size of peas, but more remarkably so in the males…about two or three hundred miles to the North West of Churchill, I have seen them in as large flocks as the Common Wavey, or Snow Goose. The flesh of this bird is exceedingly delicate; but they are so small, that when I was on my journey to the North I eat two of them one night for supper. I do not find this bird described by my worthy friend Mr. Pennant in his Arctic Zoology. Probably a specimen of it was not sent home [4]

Despite the fact that Hearne had shot (and eaten!) this bird, there were no specimens, and no attempt to give it a scientific name for quite some time. In 1859, Robert Kennicott sent a head, wings, tail and head of a Ross’s Goose plus a nearly complete skin (that he had obtained from Bernard Ross) to the Smithsonian [5]. The species was then formally described in 1861 as Chen rossii by John Cassin, ornithologist at the Philadelphia Academy of Sciences. I don’t really have a problem with Bernard Ross being recognized in this bird’s scientific name, for he did collect the first specimen to be preserved in a museum. But it would be fitting for the common name to be Hearne’s Goose, as Hearne actually was the first to recognize it as distinctive, and he made huge contributions to Arctic ornithology.

Ross’s Gull (Nelson 1887)

The Ross’s Gull, on the other hand, is named after James Clark Ross, but even that troubles me a bit. Ross (or a member of his party) collected the first specimens of this beautiful gull in Foxe Basin off the east coast of Nunavut’s Melville Peninsula in June 1823. Ross really did little more than send the specimens of two odd-looking gulls back to England. William Macgillivray mentioned these specimens in a 1824 in a footnote to a paper on gulls, where he says that the name Larus roseus is: “The name given pro tempore to a new species of gull, discovered by the Arctic expedition, but which is to receive its proper designation from Dr Richardson.” [6]  The ‘Dr Richardson’ that he refers to here is the great Arctic naturalist John Richardson who identified the bird as a distinct species and described it—as the Cuneate-tailed Gull Larus Rossii—in 1824 in his appendix to Parry’s journal of his second voyage.  Richardson, rather than Ross, should really have been immortalized in this species’ common name. Richardson loses doubly on this one because the rules of zoological nomenclature require that Macgillivray be listed after the scientific name because he was the first to publish—by only a few months— that name even though he was just vaguely referring to Richardson. Moreover, Macgillivray referred to the bird as Larus roseus and not Larus Rossii that Richardson seem to prefer.

RTstampThere is one other bird named Ross—the Lady Ross’s Turaco—which has, like the other Ross’s birds, what seems to me to be an inappropriate common name. Lady Eliza Solomon Ross was the wife of Major General Sir Patrick Ross [7], the governor of St Helena from 1846-1850. St Helena is a tiny tropical island in the mid-Atlantic about 2000 km west of Namibia. It became a British Crown Colony in 1836 after being ‘owned’ by the East India Company for more than 150 years. The St Helena Rosses were not, as far as I can tell. closely related to the James, John or Bernard Ross mentioned above.

Lady Ross must have kept a small menagerie or aviary on St Helena that included a turaco brought there from the west coast of Africa [8]. On a visit to England she gave two feathers and a drawing (by a Lieutenant J. R. Stack) of her bird to John Gould, presumably wondering what species it was. Based on that material, Gould described this individual as a new species at a meeting of the Zoological Society of London in 1851 [9].

In my opinion, either John Gould—or nobody—should have been immortalized in the common name of that turaco, and certainly not Lady Ross.

The naming of birds has a long and checkered—and interesting—history that continues even today. I see, for example, that the AOS checklist committee is currently wrestling with some proposed changes to the common names of North American birds, to right some perceived wrongs (Gray Jay to Canada Jay, and Rock Pigeon to Rock Dove), harmonize names between Europe and the Americas (Common Moorhen to Eurasian Moorhen, and Common Gallinule to American Moorhen), and to indicate the correct taxonomic affinities (Red-breasted Blackbird to Red-breasted Meadowlark).

It is perhaps rather ironic that four of the ornithologists directly involved in what I consider to be the misnaming of the three Ross’s birds have all been immortalized—and rightly so—in the common names of other birds—Cassin’s Auklet, Townsend’s Warbler, Gouldian Finch, Macgillivray’s Warbler, for example. While I consider the Ross’s birds to be misnamed—or at least inappropriately named—I would not suggest changing those names as they are well established. The origins of those common names also provides an interesting window on the state of ornithology in the 1800s. As Macgillivray said in the paper where he first listed the scientific name of the Ross’s Gull: “With regard to the outcry against change of names, I have only to observe, that names, as well as descriptions, must continue to fluctuate until they be rendered of such a nature as to be harmonized with common sense and sound judgment.” [10]


  • Baird SF, Brewer TM, Ridgway R (1884) The Water Birds of North America. Boston: Little, Brown & Co.

  • Cassin J ( 1861). Permission being given, Mr. Cassin made the following communication in reference to a new species of Goose from Arctic America. Proceedings of the Academy of Natural Sciences of Philadelphia 13: 72-72

  • Grant CHB (1915) On a collection of birds from British East Africa aid Uganda, presented to the British Museum by Capt. G. P. Cosens.-Part 111. Colii-Pici.  Ibis 57: 400-473
  • Hearne S (1795) A Journey From Prince of Wales’s Fort in Hudson’s Bay to the Northern Ocean. London: Strahan and Cadell.
  • Macgillivray W (1824) Descriptions, characters, and synonyms of the different species of the genus Larus, with critical and explanatory remarks. Memoirs of the Wernerian Natural History Society 5: 247-276
  • Nelson EW (1887) Report upon Natural History Collections Made in Alaska Between the Years 1877 and 1884. Washington, DC: Government Printing Office
  • Parry WE (1824) Journal of a second voyage for the discovery of a northwest passage from the Atlantic to the Pacific; performed in the years 1821-22-23, in His Majesty’s ships Fury and Hecla, under the orders of Captain William Edward Parry. London: John Murray.

  • Pennant T (1784) Arctic Zoology, 2 vols. London: Henry Hughs.

  • Ross BR (1862) List of mammals, birds, and eggs observed on the Mackenzie River District, with notices. Canadian Naturalist 7:137-155

  • Swainson C (1885) Provincial names and folk lore of British birds. London: English Dialect Society.

  • Whitman CH (1898) The birds of Old English literature. The Journal of [English and] Germanic Philology 2:149–198.


  1.  old books about bird names: see, for example, Swainson (1885) and Whitman (1898)
  2. North-Western Territory and Rupert’s Land: Ross was employed from 1843-1862 at the Hudson’s Bay Company trading posts at Norway House and York Factory in present-day Manitoba, as well as Fort Simpson, Fort Norman, and Fort Resolution in the present-day Northwest Territories
  3. Bernard Ross publications: e.g. Ross (1862); see here for a listing
  4. quotation: from Hearne 1795, page 412; Hearne wrote that passage long after he returned to England and was presumably surprised to learn that there were no specimens in Pennant’s book, published in 1784; Hearne died in 1793 and his Journey was published posthumously
  5. Ross’s Goose specimens: Kennicott also collected “a large number of individuals of this species” (Baird et al. 1884, page 446)) at Fort Resolution in 1860, and I am surprised that Cassin did not honour him in the scientific name of this species
  6. quotation from footnote on Ross’s Gull: Macgillivray 1824, page 249
  7. Lady Eliza Solomon Ross : some sources incorrectly identify this Lady Ross as Sir James Ross’s wife.
  8. west coast of Africa: possibly Angola, see Grant 1915 page 413
  9. turaco described by Gould: presumably Lady Ross later gave the turaco to Gould because it is now a specimen in the BM(NH) “the type, which is a worn and faded caged bird” (Grant 1915, page 413)
  10. quotation on changing bird names: Macgillivray 1824, page 276

A Century Ago

BY: Bob Montgomerie, Queen’s University | 12 March 2018

As you might suspect, I find the history of ornithology in particular—and the history of science in general—pretty interesting. But even I am not sure why.

RuseIn high school, history was my least favourite subject, taught by Dr A. S. H. Hill—our only teacher with a PhD (and in Political Science)—who insisted on having us memorize long strings of dates and events, aided by a rather pedantic textbook called The Modern Era.

Then in my final undergraduate year at the University of Guelph, the Philosophy Department hired a recent PhD graduate from the University of Bristol named Michael Ruse. Ruse’s first course was called something like Philosophical Foundations of the History of Science. I had taken a few philosophy courses so this one sounded right up my alley. In 1970, Ruse was a mild-mannered, genial new professor but lacked teaching experience. Since he had just completed his degree he decided to devote the whole course to the subject of his thesis research—Charles Darwin’s Origin of Species. Only 6 of us took that class, reading and discussing each chapter in Morse Peckham’s Variorum Text, following the changes that Darwin made as he revised the first 5 editions [1]. I think the other students were bored beyond belief but I was enthralled with this intimate look into the writings of the great man (Darwin), and the insights provided by Ruse.

The demands of (largely) ornithological research kept my nascent interest in history on hold for the next 30 years. Then, largely through discussions with Tim Birkhead, that interest was rekindled at the end of the the last century, and this blog and website are one result. If the proliferation of both books and publications are any indication, I am not the only person who has taken an interest in the history of ornithology of late, with a new book on the topic appearing every few months [2].

In 1984, when The Auk had just published its 100th volume, the editor—John A. Wiens—started a series that he called 100 Years Ago in The Auk because, as he said:

Reading the writings of a century ago reveals a good deal about how our current thinking was anticipated, how far we have come, how little has changed, how ornithologists approached the study of birds then, and the like. Accordingly, this and each following issue of The Auk will contain a selected excerpt from the counterpart issue of a century ago. [3]


As Wiens promised, an excerpt from a paper that had appeared in The Auk 100 years earlier was published in most issues of the journal for the next 15 years, chosen, presumably, by successive editors Alan Brush (1985-90), Gary Schnell (1991-96) and Tom Martin (1997-99). There were none published in 2000 when The Auk had an interim editor. But when Kimberly Smith took over as Editor in Chief in 2001 he saw an opportunity to start a new series that he called 100 Years Ago in the American Ornithologists’ Union. Kim has written these excellent pieces in every issue since January 2001. He did not add his byline until 2005 at the insistence of the new editor, Spencer Sealy, who wanted Kim to get full credit for his work.


Smith18While the extracts from century old papers were interesting and useful in the late 20th century, the ready availability of old issues of The Auk (and other journals) on SORA, would make the series that Wiens began less useful today. Kim Smith’s essays, on the other hand, provide unique insights into AOU conferences, initiatives and publications, often described in the context of contemporary  life and science. In his latest piece (Jan 2018), for example, Kim tells how the AOU obtained money through the Liberty Bond Program in the USA to help society members who were serving the WWI.

In his 1984 essay inaugurating the series of excerpts from a century ago, Wiens said that:

Ornithology was a different sort of discipline then, with much that we now accept as common knowledge yet to be discovered. In some respects, however, ornithologists of a century ago had as much understanding of natural phenomena as we do today, although they expressed themselves using more eloquent prose, less jargon, and far less quantitative detail. [3]

There is some truth in what he says but our ornithological predecessors were often difficult to understand, prone to using anthropomorphisms and the jargon of the day. They were also very quantitative at times but less inclined to use statistical analyses as most of the methods we use readily (and often incorrectly) today had not yet been invented, as Ted Anderson mentioned in last week’s post.


  • Peckham M, ed. (1959) The Origin of Species: a Variorum Text. Philadelphia: Univ. of Pennsylvania Press.
  • Wiens JA (1984) 100 Years Ago in The Auk. Auk 101:203


  1. Variorum Text: see Peckham (1959); for a modern online version see here
  2. New books on history of ornithology: I will be reviewing 3 of these here in the next couple of months
  3. Quotations: both from Wiens (1984)