Callow Youth

BY: Bob Montgomerie, Queen’s University | 11 December 2017

In science, controversies often arise over complex issues when researchers approach a problem from different points of view, backgrounds, and philosophies—think, for example, of the debates over nature vs nurture, selection vs drift, group vs kin vs individual selection, gradualism vs punctuated equilibria, and mechanisms of sexual selection. As Ledyard Stebbins pointed out in 1982, the resolution of these controversies often settles somewhere in the middle because, in fact, both sides were correct to some extent [1]. Usually scientific controversies churn away towards some resolution with some degree of civility, at least in public forums. But not always…

I can still remember my first experience with a less-than-polite public argument among ornithologists. This occurred during a talk at an AOU meeting in the 1970s when two prominent systematists got into a shouting match during the question period after a graduate student’s talk. The issue was whether the approach taken by the student was correct, and the argument seemed to be between the cladists and the pheneticists.

I learned later that these clashes among systematists were commonplace in the 1960s and early 70s, during a period that was later referred to as the ‘Systematics Wars’ [2]. I was doubly surprised at such ad hominem attacks because the systematists that I knew from my years as a young volunteer and employee at the Royal Ontario Museum—L.L. Snyder, Jim Baillie, Jon Barlow, Jim Rising, and Allan Baker—seemed like the kindest of men. My own field of behavioural ecology, while often dealing in controversy, does not seem to have descended into the sort of personal attacks that characterized those arguments about systematics in the mid 1900s.

While scouring some older literature, we recently discovered [3] that the ‘Systematics Wars’ period was not the first time that avian systematists had engaged in rather nasty exchanges. During the 1830s, for example, two young  brothers—Charles Thorold Wood and Neville Wood, tried to change the rules about naming birds in a way that involved heated debate, vitriol, and ad hominem attacks in their publications.

The Wood boys were wealthy, aristocratic, well-educated, and precocious. In 1835, when he was only 18, Charles published a quirky book—The Ornithological Guide—comprising 236 pages of poetry about birds, a compendium of ornithological books, each briefly reviewed, and a catalog (list) of the birds of Britain. Not to be outdone by his older brother, Neville published two books the next year, when he turned 18—British Song Birds and Ornithologists’ Text-book. British Song Birds alone ran to 400 pages, and, though it appeared not to include any novel observations, summarized much of what was then known about each species. Notably, none of the Woods’ publications included any illustrations.

The Woods felt that the names of birds—both scientific and English—were in a chaotic state and needed rules to make them more ‘scientific’. They were right. Even though Linnaeus had proposed his binomial system almost a century earlier [4], rules for zoological nomenclature that would provide some consistent structure and process were just beginning to be proposed in the 1830s [5].

Female and male European Bullfinches, by John Gould 1837 in his Birds of Europe

The English names were more confusing and disorganized. In England, some birds were even given different English names in different parts of the country. In the 1800s, for example the European Bullfinch was variously called: Bull Flinch (Yorks), Bull Head, Bulldog, Bull Spink, Bully (Yorks), Thick Bill (Lancs), Alpe, Hoop, Hope (SW), Tope, Hoof, Cock Hoop (Hereford), Olf (E Suffolk), Nope (Staffs/Shrops), Mwope (Dorset), Mawp (Lancs), Pope (Dorset), Red Hoop (m, Dorset), Blood Olp (m, Surrey/Norfolk), Tawny (f, Somerset), Tony Hoop, Tonnihood (f, Somerset), Black Cap (Lincs), Billy Black Cap, Black Nob (Shrops), Monk, Bud Bird, Bud Finch, Bud Picker (Devon), Budding Bird (Hereford), Plum Bird, and Lum Budder (Shrops) [6].

The Woods were sure that some logic and rules for the structure of English names were needed if ornithology was to develop as a science. As Neville said in one paper ”“If the proper English generic names were applied to every bird, how greatly would the acquisition of this fascinating study be facilitated! . . . By using the names which I have given above, this [confusion] is remedied, and all becomes plain and easy to understand.” [7] Their solution, for English names, was to make every bird within a genus have the same ’surname’, just as every genus had the same first name in the Linnaean system of Latin names. Thus, for example, Neville proposed:


It was also suggested  at the time [8] that English names be cleansed of modifiers that indicate size, abundance, location, or honorific. Thus present-day names like Little Cuckoo, Common Cuckoo, Moluccan Cuckoo, and Klaas’s Cuckoo would all have to be changed in the Cuculidae alone. Fortunately, cooler heads prevailed, as Hugh Strickland argued that common names were “consecrated by usage as much as any other part of the English language”. As he noted “the science of ornithology does not suffer by this incorrect application of English names, because those familiar appellations have no real or necessary connection with science”. Despite his admonition, the English names of birds are still in a state of flux today and may always be.

The Woods’ proposals were well reasoned, if sometimes seeming to be a little bizarre today. But their suggestions were often presented in such a way that demeaned those who had different points of view. In the Preface to his Bird Song Birds, Neville says: “while I agree with my predecessors in many points, I have found much to correct, and still more to add, to the meagre and unsatisfactory accounts of most of our British Ornithologists.” Ouch!

Neville’s review of Eleazar Albin’s Natural History of Birds claimed that it was “of no use in the present day” and of Jennings Ornithologia that he ”never had the misfortune to meet with a book so full of errors . . . We should have considered such a work beneath our notice, as it is impossible it can have the smallest connection with the advancement of Ornithology”. [9] Their writings are sprinkled with derogatory epithets and harsh criticism that even today strikes me as ungentlemanly, possibly simply the result of the Woods being callow youth [10].

Possibly because they gained few converts, and appear not have been well liked, both Charles and Neville drifted off to other pursuits before their 35th birthdays, never again to write about birds or to heap opprobrium on their fellow ornithologists. Charles published his last paper on birds (again on nomenclature) when he was 19 and then seems to have vanished from the public record; Neville trained as a doctor, and at the age of 26 moved to London where he practiced the new medical system called ‘homeopathy’ for the rest of his life. At least if he had stayed with birds he would have done something potentially useful.

Stebbins ended his 1982 address to the American Association for the Advancement of Science [11] by saying that: “My final hope is that evolutionists having different backgrounds and viewpoints will reduce their rivalry and collaborate increasingly in zealous research toward finding answers to these and other questions of major significance.” This was a noble sentiment but ignored those ineluctable human (or at least male) foibles of ego, ambition, and competitiveness. I would argue that controversies are are valuable part of scientific progress, and I have certainly enjoyed watching the systematics wars from the sidelines. We should always try to maintain civility but the controversies that arise from different points of view often move a field forward faster and more profitably than would the absence of skepticism about published research.


  • Albin E (1731) A Natural History of Birds : Illustrated With a Hundred and One Copper Plates, Curiously Engraven From the Life. Vol I. London: Printed for the author and sold by William Innys in St. Paul’s Church yard, John Clarke under the Royal-Exchange, Cornhill, and John Brindley at the King’s Arms in New Bond-Street. <available here>

  • Anonymous “N. F.” (1835) Remarks on vernacular and scientific ornithological literature. The Analyst 2: 305–307.

  • Birkhead TR, Montgomerie R (2016) A vile passion for altering names: the contributions of Charles Thorold Wood jun. and Neville Wood to ornithology in the 1830s. Archives of Natural History 43:221–236.

  • Greenoak F (1997) British Birds: their folklore, names, and literature. London: Christopher Helm.

  • Jennings J (1829) Ornithologia, or, the Birds: A Poem, in Two Parts : With an Introduction to Their Natural History; and Copious Notes. London: Sherwood, Gilbert, and Piper. <available here>

  • Stebbins GL. (1982a). Modal themes: a new framework for evolutionary synthesis. in Milkman R (Ed.), Perspectives on Evolution (pp. 12-14). Sunderland, MA: Sinauer Associates.
  • Stebbins GL (1982b) Perspectives in evolutionary theory. Evolution 36:1109–1118.
  • Sterner B, Lidgard S (2017) Moving past the Systematics Wars. Journal of the History of Biology
  • Wood CT (1835) The ornithological guide: in which are discussed several interesting points in ornithology. London Whittaker. <available here>
  • Wood N (1836a) British song birds: being popular descriptions and anecdotes of the choristers of the groves. London: John W. Parker. <available here>
  • WOOD N (1836b) The ornithologists’ text-book: being reviews of ornithological works on various topics of interest. London: John W. Parker. <available here>


  1. Stebbins (1982a) argued that modal themes were often the correct resolution of these conflicts, as has proven to be more-or-less the case in all the controversies listed here, with the notable exception that group selection is still controversial (and mistaken in my opinion)
  2. see, for example, Sterner and Lidgard (2017) for an overview
  3. Tim Birkhead made this discovery while researching for the project on Francis Willughby that he wrote here about a few weeks ago
  4. Linnaeus Systema Naturae had built on Willughby and Ray’s attempt at organizing and logically giving each species a scientific name (Ray (1678), but in the subsequent century there were many attempts to apply Linnaeus system to the birds with each author claiming authority and none of today’s rules about priority, coordination or homonymy.
  5. The International Zoological Congresses of 1889 and 1892 saw the first discussions about some universal rules in zoology but the International Rules on Zoological Nomenclature were not published until 1905.  Earlier, the ornithologist Hugh Strickland had published a set of 22 rules for the formulation of scientific names that formed some of the basis for this later code.
  6. the sex that was so named,  and the county or general area where each name was used, are shown in brackets, from Greenoak (1997).
  7. quotation from Wood 1835a: 238
  8. this was proposed in a paper in The Analyst signed simply N.F. (Anonymous 1835), who I think may well have been Neville Wood based on what N.F. says and the style of writing.
  9. see Birkhead and Montgomerie (2016) for some other gems. The paper is behind a paywall but send me an email if you cannot get it and would like a copy.
  10. ‘callow’ seems to me to be a particularly good adjective to describe these young men, as it refers to someone who is “inexperienced and immature”,  but in the 17th century the term was applied to birds, and meant ‘without feathers’ or immature and not ready to fly.
  11. presented on 7 Jan 1982, at a AAAS symposium marking the 100th anniversary of the death of Charles Darwin; quotation from Stebbins (1982b)

Pigeon Coup

BY: Bob Montgomerie, Queen’s University | 27 November 2017

When I was a young teenager I spent my Saturday mornings during the school year at the Royal Ontario Museum (ROM). I was there to attend the weekly meeting of the Toronto Junior Field Naturalists’ Club, but often stayed afterward to explore the public galleries. I particularly loved the dioramas of birds and mammals as they took me to distant places and bygone times that I could only dream or read about. In those days, there were virtually no nature documentaries on TV and precious few in the theatres [1].

Of all those superb dioramas, my favourite was the Passenger Pigeon, showing immense flocks (painted) descending into a forest clearing scattered with pigeons (mounted specimens) foraging on the acorns:

The scene reproduced depicts an April morning in the 1860s near Forks of Credit, Ontario…The visitor inspecting the exhibit should imagine himself standing at the edge of an old beech-maple forest overlooking the pioneer’s clearing. The scene is as we might have found it in the 1860s. The great pigeon flight is underway and will perhaps continue throughout the day. [2]

That diorama was opened to the public in 1935, the brainchild of Lester L. Snyder, the curator of birds, and constructed and painted by E. B. S. Logier who had joined the museum as illustrator in 1915. My memories of those dioramas came flooding back a couple of weeks ago when I saw the names Mark Peck and Allan Baker [3]—both from the ROM—among the authors of a new paper out of Berth Shapiro’s lab (UC Santa Cruz) on Passenger Pigeons published in Science, but more on that in a minute.

13662-Passenger Pigeon-retouched
Painted backdrop of the Passenger Pigeon diorama at the ROM (1935-1981)

Standing in front of that diorama I can remember thinking that a bird that had once been that abundant could not possibly be extinct. After all, Peterson’s Field Guide still illustrated them (in  a head and shoulders vignette on p 181 of my copy) so maybe he thought the bird might still be seen. My teen birding buddies and I spent many an afternoon naively scouring flocks of Mourning Doves just in case. After all, bison were also once extremely abundant, and hunted relentlessly, but were still round, albeit in small numbers.

I was also heartened by the fact that even if the species was really extinct, there must be thousands of specimens in museum collections that could be used for further study, since the ROM alone appeared to have so many that they could fill a diorama with mounted specimens alone. When I mentioned this to my friend and mentor Jim Baillie, assistant curator of birds at the ROM, he just laughed and told me there were only about 1500 specimens worldwide, of a species that once numbered in the billions. The reason for this wealth of specimens at the ROM, he said, was that they had been the beneficiaries of what he considered to be a major coup, when a local naturalist, musician and businessman [4], Paul Hahn, had decided—shortly after the Passenger Pigeon went extinct in 1914—to donate to the ROM as many specimens as he could locate, as a way “to ensure that future generations would know at least how handsome a bird it was.” [5]

Paul Hahn

Hahn was born in Germany in 1875 but moved to Toronto with his family in 1898. In 1902 he saw his first Passenger Pigeon, a mounted specimen in a farmhouse north of the city and decided then to “set about gathering as many as possible of the mounted birds scattered around the country, both for the sake of future students and with the intention of preserving at least some specimens of a bird that would probably soon be extinct.” [5]. He presented his first specimen to the ROM in 1918 and had donated 70 by the time he died in 1962.

As a result of Hahn’s generosity, the ROM had 124 Passenger Pigeon skins and mounts by 1962, more than any other collection worldwide. I know this because in 1957 Mr Hahn started compiling a list of all the specimens of 7 extinct (our nearly so) bird species [6] held in museums and private collections around the world. Hahn died before his list could be published but Baillie took up the task, seeing it through to print in 1963 as a book Where is that Vanished Bird? That book lists every specimen (including skeletons) known to Hahn [7], its date and place of collection, its sex, the collector, and the current collection in which it was held.

One of the Passenger Pigeon mounts at the Royal Ontario Museum

The first Passenger Pigeon specimen whose collection date was known was a male taken in the Carolinas in about 1810, housed with one other specimen (a female, from Georgia collected in 1821) in the Zooligische Museum in Berlin. The Naumann Museum in Köthen that Tim Birkhead wrote about last week also had a male, collected in about 1830 (locality unknown).

The recent Science paper made good use of the ROM collection of Passenger Pigeons, analyzing the DNA extracted from the toe pads of 84 specimens, 63 of which were from the ROM. Analyzing both nuclear and mitochondrial genes, the researchers confirmed that the Passenger Pigeon had surprisingly low genetic diversity. This low diversity is unexpected because large populations are predicted from theory to be genetically diverse, and you don’t get larger bird populations than those of the Passenger Pigeon. To explain this loss of diversity, the authors argued that it was driven by high rates of dispersal and adaptive evolution that removed harmful mutations. Such low diversity would have made the species particularly susceptible to disease or environmental change, two factors that might have doomed the species once populations had been decimated by hunting. This study also concluded, based on some sophisticated genomic analyses, that Passenger Pigeon populations had probably persisted at extremely high numbers for 20,00 years or more before the 1800s [8].

The Passenger Pigeon diorama at the ROM was dismantled in 1981, in part because it was showing its age, but also because the age of dioramas was over, replaced in part by the ubiquitous nature shows in TV. That saddens me but I am more than ever convinced that clubs for young field naturalists, and museums that store and preserve specimens, deserve our unending support.


  • Hahn P (1963) Where is that Vanished Bird? Toronto: University of Toronto Press.

  • Hung C-M, Shaner P-JL, Zink RM, Liu W-C, Chu T-C, Hiuang W-S et al. (2014) Drastic population fluctuations explain the rapid extinction of the passenger pigeon. Proceedings of the National Academy of Scienes USA 111:10636–10641.

  • Murray GGR, Soares AER, Novac BJ, Schaefer NK, Cahill JA, Baker AJ et al. (2017) Natural selection shaped the rise and fall of passenger pigeon genomic diversity. Science 358:951–954.


1. The first nature documentaries on TV were a series called Fur and Feathers on Canadian Broadcasting Corporation (CBC) channel in 1955-56, in black and white (of course). By 1960, Disney had produced 14 movies in its True-Life Adventures series, including the The Living Desert, The Secrets of Life, African Lion, and White Wilderness all of which enthralled my naturalist friends and I when they played at our local theatre.

2. Text from the ROM’s Passenger Pigeon diorama, courtesy Mark Peck, 22 Nov 2017.

3. Mark Peck is Ornithology Technician at the ROM, where Allan Baker (1943-2014) worked for 42 years as both a curator of birds and eventually head of their Department of Natural History.

4. Hahn was an accomplished cellist who played with the Toronto Symphony Orchestra. He also founded Paul Hahn Pianos in Toronto in 1913, a company that is still in business today.

5. Quotations from Hahn 1963:1.

6. As of 1962: skins and mounts of 1532 Passenger Pigeons, 365 Eskimo Curlews, 78 Great Auks, 720 Carolina Parakeets, 413 Ivory-billed Woodpeckers, 54 Labrador Ducks and 309 Whooping Cranes (Hahn 1963).

7. Beginning in 1957 he sent out questionnaires to people and museums that he thought might know or know about those specimens. He got more than 1000 response.

8. Based on DNA samples from only 3 Passenger Pigeons, Hung et al. (2014) performed a different genomic analysis and concluded that population sizes had fluctuated dramatically—only occasionally reaching numbers in the billions—thereby increasing its risk of extinction during population lows. Evaluating the conclusions of these two studies is above my pay grade but I expect that both labs will argue that their analyses are correct.

IMAGES: ROM photos by Brian Boyle, courtesy of Mark Peck (both at the ROM); Paul Hahn from the Paul Hahn & Co. website at

Birds and Revolutions

BY: Bob Montgomerie, Queen’s University | 28 Aug 2017

This month [see footnote 1] marks the anniversary of the famous 1858 Darwin-Wallace publication on natural selection published in the Journal of the Proceedings of the Linnean Society of London, Zoology (PDF here). [Note that it was read at a meeting of the Linnean Society on 1 July and published a mere 7 weeks later!] This joint paper ushered in what is arguably the greatest scientific and cultural revolution in human history, a fine example in support of Thomas Kuhn’s (1962) argument that scientific revolutions are driven by ideas. Last week I presented, on this blog, some evidence that tools also fuel revolutions in science.


How did the study of birds figure into Darwin and Wallace’s arguments? We know from Darwin’s many books, and much of Wallace’s writing, that both men studied birds and used examples from birds to illustrate and support many—if not most—of their ideas. This 1858 paper marks the beginning of the association between birds and natural selection. We cannot argue that the study of birds was in any direct way responsible for this scientific revolution but there can be no doubt that this revolution forever influenced how we study birds.

In this publication, Darwin first uses birds as an example of Malthus’s ideas about population:

Suppose in a certain spot there are eight pairs of birds, and that only four pairs of them annually (including double hatches) rear only four young, and that these go on rearing their young at the same rate, then at the end of seven years (a short life, excluding violent deaths, for any bird) there will be 2048 birds, instead of the original sixteen. As this increase is quite impossible, we must conclude either that birds do not rear nearly half their young, or that the average life of a bird is, from accident, not nearly seven years. Both checks probably concur. The same kind of calculation applied to all plants and animals affords results more or less striking, but in very few instances more striking than in man

He then introduces his arguments about sexual selection that he will work on for another 13 years before publishing in 1871 in Descent of Man.

These struggles are generally decided by the law of battle, but in the case of birds, apparently, by the charms of their song, by their beauty or their power of courtship, as in the dancing rock-thrush of Guiana. The most vigorous and healthy males, implying perfect adaptation, must generally gain the victory in their contests. This kind of selection, however, is less rigorous than the other; it does not require the death of the less successful, but gives to them fewer descendants. The struggle falls, moreover, at a time of year when food is generally abundant, and perhaps the effect chiefly produced would be the modification of the secondary sexual characters, which are not related to the power of obtaining food, or to defence from enemies, but to fighting with or rivalling other males.

Wallace also uses birds to illustrate Malthus’s ideas, though, unlike Darwin, he never refers directly to Malthus in his essay:

For example, our own observation must convince us that birds do not go on increasing every year in a geometrical ratio, as they would do, were there not some powerful check to their natural increase. Very few birds produce less than two young ones each year, while many have six, eight, or ten; four will certainly be below the average; and if we suppose that each pair produce young only four times in their life, that will also be below the average, supposing them not to die either by violence or want of food. Yet at this rate how tremendous would be the increase in a few years from a single pair! A simple calculation will show that in fifteen years each pair of birds would have increased to nearly ten millions! whereas we have no reason to believe that the number of the birds of any country increases at all in fifteen or in one hundred and fifty years. With such powers of increase the population must have reached its limits, and have become stationary, in a very few years after the origin of each species. It is evident, therefore, that each year an immense number of birds must perish—as many in fact as are born; and as on the lowest calculation the progeny are each year twice as numerous as their parents, it follows that, whatever be the average number of individuals existing in any given country, twice that number must perish annually,—a striking result, but one which seems at least highly probable, and is perhaps under rather than over the truth.

Finally, I particularly like the following passage from Wallace’s contribution to this publication, where he makes an ecological argument for the abundance of the passenger pigeon:

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 differences 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. 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.

…and continues with ideas about the adaptive significance of migration and factors that influence the relative abundance of species:

In no other birds are these peculiar circumstances so strikingly combined. Either their food is more liable to failure, or they have not sufficient power of wing to search for it over an extensive area, or during some season of the year it becomes very scarce, and less wholesome substitutes have to be found; and thus, though more fertile in offspring, they can never increase beyond the supply of food in the least favourable seasons. Many birds can only exist by migrating, when their food becomes scarce, to regions possessing a milder, or at least a different climate, though, as these migrating birds are seldom excessively abundant, it is evident that the countries they visit are still deficient in a constant and abundant supply of wholesome food. 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. Why, as a general rule, are aquatic, and especially sea birds, very numerous in individuals? Not because they are more prolific than others, generally the contrary; but because their food never fails, the sea-shores and river-banks daily swarming with a fresh supply of small mollusca and crustacea.

Most biologists have read little of what Darwin and Wallace wrote, in part because they think it’s out of date (it’s not) and both convoluted and a bit dry to read (it is, but not uninteresting). If you read nothing else written by these two men, this initial publication, with essays by both of them, is well worth the few minutes invested to read and, particularly, to think about the revolution they started 159 years ago.


Darwin CR (1871) The Descent of Man and Selection in Relation to Sex. John Murray, London.

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

Kuhn TS (1962) The Structure of Scientific Revolutions. University of Chicago Press, Chicago.


  1. There seems to be some confusion about the actual date. Wikipedia says “the papers appeared in print on 20 August 1858” as does the Royal Society’s twitter feed, which (strangely) links the reader to Wikipedia (see below). Darwin Online, however, says “August 30th, when it appeared in print”.