Bird Paper Two

BY: Bob Montgomerie, Queen’s University | 6 May 2019

A few months ago (10 December 2018), I wrote about the first paper ever published about birds (here)—a description of a hummingbird from Barbados, published by the botanist Nehemiah Grew in May 1693. This publication was in the Royal Society’s Philosophical Transactions, and did not appear until 28 years after scientific publishing began, in 1665. Ornithology did not really become a ‘science’ in England until Ray’s The Ornithology of Francis Willughby was published in English in 1678 [1] but it still took another 15 years before a scientific paper on birds appeared.

Grew also published the second-ever paper about birds when he followed up on his first publication in the July/August 1693 issue of Philosophical Transactions. That second paper is short enough that I can reproduce it here in its entirety:


In that second paper he reports that the observations that he reported on earlier that year were actually made by a Mr Hamersly, and he wonders if the birds really do subsist on nectar, or are actually eating insects on the bottoms of corollas. He must have known that none of the English birds that he would have been familiar with subsisted on nectar alone so it would have seemed anomalous to him that any bird could. He wisely suggests opening up a hummingbird to see if they have different sorts of guts from other birds.

That was a great suggestion—and would have been a revelation—as the stomach of a hummingbird is a specialized structure. At least in the few species whose stomachs have been studied, food enters the hummingbird’s proventriculus (stomach) very close to and on the same plane as where it exits into the duodenum from the ventriculus. Thus the liquid nectar takes a shortcut to the small intestine without passing into the ventriculus. The ventriculus is used to crush insects that the bird eats but really has nothing to offer the nectar. Nutrients are absorbed in the small intestine so the hummingbird digestive systems gets digested nectar to the site of absorption as quickly as possible.

Digestive system of the chicken
Female Broad-tailed Hummingbird at nest

When I was analyzing data for my PhD thesis (on hummingbirds), I lived for a month in a VW camper in the mountains north of Tucson, Arizona. A female Broad-tailed Hummingbird came into our camper regularly to inspect the red plastic plates by the sink. This surprised me because I saw no obvious hummingbird flowers during my frequent walks in the woods. One day, I followed the female back to her nest and often saw her foraging on insects.

We surveyed the woods for 500 m in each of the eight cardinal directions from the nest and found no flowers whatsoever. We also watched the bird during 19 foraging bouts away from the nest and only saw her catching aerial insects and taking prey from spiders’ webs. As far as we were aware at the time (and still), this was the first evidence that hummingbirds could subsist solely on arthropods for a while, and it makes sense, from a nutritional point of view, that they would have to feed animal protein to their nestlings, as otherwise they could not grow.

Hummingbirds continue to be a fascinating group to study but Grew was not the first to publish something about them. That honour goes to the French explorer Jean de Léry who spent 10 months on the coast of Brazil in 1557. de Léry published about his adventures in 1578, with a chapter on the birds he saw, including “une singuliere merveille, et chef-d’oeuvre de petitesse, il n’en faut pas omettre un que les sauvages nomment Gonambuch, de plumage blanchastre et luisant, lequel combien qu’il n’ait pas le corps plus gros qu’un frelon, ou qu’un cerf-volant, triomphe neantmoins de chanter” [2], in other words, a hummingbird.

de Léry’s description could hardly be called ‘scientific’, so the first description of a hummingbird—in fact of 9 species—that is both accurate and scientifically interesting was not published until almost a century later, in 1648. These descriptions appear in Georg Marcgrave‘s section of Historia naturalis Brasiliae. Marcgrave’s original text is in Latin, and was later translated into French and English [3]. I will post a full account of Macgrave’s findings later this year. In the meantime, here is a picture (from Marcgrave) of one of the species he describes:

Guainumbi (hummingbird) in Marcgrave (1648)


  • Grew N (1693a) The description of the American tomineius, or hummingbird. Philosophical Transactions 17: 760-761
  • Grew N (1693b) A query put by Dr. N. Grew, concerning the food of the Humming Bird; occasioned by the description of it in the transactions. Numb. 200. Philosophical Transactions 17: 815
  • Klasing K (1998). Comparative Avian Nutrition. New York: CAB International.
  • de Léry J (1578) Histoire d’un voyage fait en la terre du Bresil, autrement dite Amerique. Contenant la navigation, et choses remarquables, veuës sur mer par l’aucteur. Le comportement de Villegagnon en ce pays la. Les mœurs et façons de vivre estranges des Sauvages Ameriquains : avec un colloque de leur langage. Ensemble la description de plusieurs Animaux, Arbres, Herbes, et autres choses singulieres, et du tout inconnues pardeçà: dont on verra les sommaires des chapitres au commencement du livre. Le tout recueilli sur les lieux par Jean de Lery, natif de la Margelle, terre de sainct Sene, au Duché de Bourgongne. La Rochelle ou Genève: Antoine Chuppin. [1611 edition available here]
  • de Léry J (1990) History of a Voyage to the Land of Brazil. Translation and introduction by Janet Whatley. Berkeley: University of California Press
  • López-Calleja MV, Fernández MJ, Bozinovic F (2003) The integration of energy and nitrogen balance in the hummingbird Sephanoides sephaniodes. Journal of Experimental Biology 206:3349–3359.
  • Marcgrave G (1648) Historiae Naturalis Brasiliae, Liber Quintus, Qui agit de Avibus. in Piso et al. (1648)
  • Montgomerie RD, Redsell CA (1980) A nesting hummingbird feeding solely on arthropods. Condor 82:463–464.
  • Piso W, Hackius F, Laet JD, Marggraf G, Lud. E (1648) Historia naturalis Brasiliaeauspicio et beneficio illustriss. I. Mauriti Com. Nassau illius provinciae et maris summi praefecti adornata: in qua non tantum plantae et animalia, sed et indigenarum morbi, ingenia et mores describuntur et iconibus supra quingentas illustrantur. Leiden: Lugdnum Batavorum, Apud Franciscum Hackium, et Amstelodami apud L. Elzevirium. [available here]
  • Ray J (1678) The Ornithology of Francis Willughby. London: John Martyn.


  1. published in English in 1678: it was first published in Latin in 1676
  2. quotation from de Léry: (1578) which I translate as “But for a singular marvel, and masterpiece of smallness, I must not omit one that the savages call Gonambuch, with whitish and shiny plumage, which has a body no bigger than a hornet, or a beetle, nevertheless triumphs to sing”. Whatley’s (1990) translation of this passage is slightly different but the details are the same.
  3. translated into French and English: while I studied both French and latin for 5 years at school, my Latin is rustier from lack of use (!). I have not located the French and English translations in a library or on the web.

IMAGES: chicken digestive system from Wikipedia; Grew paper from Biodiversity heritage Library; Broad-tailed hummingbird photo by Bill Ratcliff from Wikimedia Commons

Mr. Cairngorms

BY: Bob Montgomerie, Queen’s University | 18 February 2019

In 2013, while compiling information for a chapter on the contributions of ornithology to evolutionary biology, I carried on a lively correspondence, by email, with Adam Watson. Watson was a renowned Scottish ecologist, naturalist and conservationist who had worked with Vero Wynne-Edwards, a staunch promoter of evolution by group selection.  We have all of Wynne-Edwards archives here at Queen’s University [1] but the man was exceptionally careful not to record (or preserve) any personal opinions in his letters or field notes. I wrote to Watson to learn more about Wynne’s interactions with David Lack, the leading evolutionary ecologist of the day, who saw no need for anything different from the sort of individual selection described by Darwin.

Wynne-Edwards and Watson had, for example, taken David Lack on a walk in the Cairngorms in 1968 at a time when the two men were having a fierce debate about group selection [2]. But, as Watson wrote to me, their meeting was extremely amicable and group selection was, as far as he could remember, never mentioned. Lack impressed everyone by spotting a rare bird and a rare plant [3], and the day turned out to be a pleasant hike in the mountains with a focus on natural history.

Adam Watson banding a ptarmigan chick

Watson was on that day-hike with Lack and Wynne-Edwards, not only because he worked with Wynne, but also because he was an expert on the Cairngorms, three plateaus of wild natural beauty in the eastern highlands of Scotland, dotted with the blunted fists of mountains rising above the plateaus. Five of those mountains are among the 6 highest peaks in Scotland and snow can fall at any time of the year. I have never been to that part of Scotland but descriptions of both the landscape and the birds remind me of High Arctic Canada, with breeding Snow Buntings and Rock Ptarmigan, and even the occasional Lapland Longspur and Snowy Owl. Watson loved it there and spent much of his life exploring and studying the region. He certainly earned the local moniker of ‘Mr Cairngorms’.

I first encountered Watson’s research in the 1970s in a graduate course at UBC in ecology where we read his classic papers with Jenkins and Moss on the causes of red grouse cycles in Scotland. These Scottish biologists were clearly in the Elton/Chitty/Krebs school of ecology, wherein they attributed population fluctuations to extrinsic factors like food supply and predation.

Red Grouse numbers on one study area in Scotland, 1957-1961

My second encounter with Watson’s work began in the 1980s when I started studying Rock Ptarmigan in the Canadian High Arctic. Watson had done his own PhD in 1956 on Rock Ptarmigan in Scotland. He continued to study that species at least until 1964 but then turned his attention to the more widespread and economically important Red Grouse. We were interested in behavioural ecology, specifically sexual selection with respect to combs and plumages, so it was important for us to keep the basic ecology of ptarmigan (of which the Red Grouse is one) in mind. We were also inspired by the experimental approach that Watson and colleagues took to addressing questions of interest.

  During the 1980s, Peter Hudson, Andy Dobson and their colleagues discovered a role for parasites in these population fluctuations. The result was a fierce conflict between Watson and the others about the controlling factors in population regulation. Thirty 30 years later, now that the dust has settled a bit, I expect that both camps had useful data and arguments to bring to bear on population cycles.

Most recently, when writing about the history of ornithology in Nunavut, I read Watson’s book [4] and papers from his summer on Baffin Island in 1953. In 1945, when he was only 14, Watson met Wynne-Edwards when Wynne moved to Aberdeen as Regius Professor. Wynne’s first academic position was at McGill University where he lectured from 1929 until the start of WWII. With their shared interest in natural history, Wynne and Watson often hiked in the mountains of northeastern Scotland.

Watson did his honours thesis at Aberdeen University studying ptarmigan under Wynne’s supervision, then in 1952 began his PhD continuing his ptarmigan research. At Wynne’s encouragement, Watson applied for and won a Carnegie Arctic Scholarship to attend McGill University for a year, associated with the nearby Arctic Institute of North America. It was there that the Director, Pat Baird, invited Watson to be zoologist on a 13-man expedition [5] to Baffin Island in 1953.

Watson studying Snowy Owls (sketch by James Houston [6]
The purpose of that expedition was to study the geology, glaciers, zoology and botany of an area called the Penny Highlands on the Cumberland Peninsula on the southeast coast of Baffin. They departed Montreal on 12 May and returned 4 months later. Watson conducted probably the first reasonably accurate census of the densities of tundra-breeding birds, and did a comprehensive study of Snowy Owl breeding biology and ecology. In his studies of both the owls on Baffin and the ptarmigan in Scotland, Watson was a pioneer in the field that we now call behavioural ecology.

WatsonVCWEWatson was clearly thrilled with his experiences on that expedition: It was a very fine summer trip. Being on that expedition with my expedition colleagues and Inuit companions, and studying Arctic wildlife among the finest mountains I had ever seen, were a rare treat and a highlight of my life. Perhaps they were the highlight.[7].

One clear attraction was that that region in particular, and Baffin Island in general, had rarely been explored by scientists. Only two ornithologists had so far done any work on Baffin: Bernhard Hantzsch on an ill-fated expedition to cross the island in 1909-1911, and Dewy Soper looking for the breeding grounds of the blue goose in the 1920s. Watson thus did not really know what to expect on the Penny Highlands so he did both general surveys and focused studies.

Toward the end of the summer they were joined by Wynne-Edwards who had been studying seabirds at Cape Searle at the tip of the Cumberland Peninsula. In preparation for their trip back to civilization everyone got their hair cut. I expect that Watson really treasured the photo above where his PhD supervisor was cutting his hair in one of the most beautiful outdoor barbershops in the world.

Adam Watson died [8] on 23 January 2019. He was 88 years old. I never had the privilege of meeting him, but I feel honoured to have known him even just by email. Someday I will visit the Cairngorms to pay tribute to his tireless efforts to preserve that interesting part of Scotland for both the birds and hares, and for people like him that love wild places.



  • Birkhead TR, Wimpenny J, Montgomerie R (2014) Ten Thousand Birds: Ornithology since Darwin. Princeton, NJ: Princeton University Press.

  • Hudson P, Tapper S (1979) Grouse populations—do they cycle? Annual Review of Game Conservancy 11:17–23.
  • Hudson PJ, Dobson AP,  Newborn D (1985) Cyclic and non-cyclic populations of red grouse: a role for parasitism? In: Ecology and genetics of host-parasite interactions (Ed by Rollinson D, Anderson RM). Pp 79-89. London: Academic Press. London.
  • Hudson PJ (1986) The effects of parasitic infections on the population fluctuations of red grouse in the north of England. In: Proceedings of the Third International Grouse Symposium (ed. By Hudson PJ, Lovell TWI )
  • Jenkins D, Watson A, Miller GR (1963) Population Studies on Red Grouse, Lagopus lagopus scoticus (Lath.) in North-East Scotland. Journal of Animal Ecology 32: 317-376
  • Montgomerie RD (2018) History of ornithology in Nunavut. pages 45-69 in Richards JM, Gaston AJ, editors. Birds of Nunavut. Vancouver: UBC Press.
  • Watson A (1953) Birds in Cumberland Peninsula, Baffin Island. Canadian Field-Naturalist 71:87–109.
  • Watson A (1956) The annual cycle of rock ptarmigan. Ph.D. thesis. Aberdeen Univ., Aberdeen, Scotland.
  • Watson A (1957) The behaviour, breeding and food-ecology of the snowy owl Nyctea scandiaca. Ibis 99:419–462.

  • Watson A (1963) Bird numbers on tundra in Baffin Island. Arctic 16:101–108.

  • Watson A (1965) A population study of ptarmigan (Lagopus mutus) in Scotland. Journal of Animal Ecology 34: 135-172.
  • Watson A (2011) A Zoologist on Baffin Island 1953. Rothersthorpe, UK: Paragon Publishing.

  • Watson A, Moss R (1970) Dominance, spacing behaviour and aggression in relation to population limitation invertebrates. In A. Watson (Ed.), Animal populations in relation to their food (pp. 167-220). Blackwell Sci.
  • Watson A, Moss R (1971) Spacing as affected by territorial behaviour, habitat and nutrition in red grouse (Lagopus l. scotius). In A. H. Esser (Ed.), Behaviour and environment; the use of space by animals and men (pp. 92-111). New York and London: Plenum Press.
  • Watson A, Moss R (1977) Population ecology of red grouse. Annual Report of the Institute of Terrestrial Ecology 1977 1978:18–21.
  • Watson A, Moss R (1979) Population cycles in the Tetraonidae. Ornis Fennica 56:87–109.


  1. archives at Queen’s University: Wynne-Edwards’s grand-daughter, Kathy Wynne-Edwards, was one of my faculty colleagues here in the Biology Department when Wynne-Edwards died, and a few of his relatives also lived in Kingston so this was a logical place for his archives to be deposited. See here for more details.
  2. Debate about group selection: see Birkhead et al. 2014 pp 369-371
  3. rare plant: the Lacks were on that hike especially to look for the rare Tufted Saxifrage (Saxifraga caespitosa) which was purported to be there.They did not find it but they did find the Brook Saxifrage (Saxifraga rivularis), a new site for a species that was also quite rare [see CORRECTION below]
  4. Watson’s book: self-published in 2011, full 58 years after he went to Baffin Island this chronicle of his summer on the Canadian tundra is remarkably detailed and full of both adventure and natural history.
  5. sketch by James Houston: Houston was a Canadian artist and novelist who lived for many  years in Cape Dorset on Baffin Island; he is credited with launching the production of Inuit prints and sculpture
  6. 13-man expedition: only 12 returned as one of the glaciologists, Ben Battle, drowned and was buried on the tundra
  7. quotation: from Watson 2011 page 5
  8. Adam Watson died: see here and here

IMAGES: all photos and the drawing by Houston courtesy Adam Watson; the photos of Watson and Wynne-Edwards was taken by Hans Röthlisberger; graph modified from Figure 4 in Jenkins et al. (1963) with the addition of Red Grouse illustration from Birds of the World Online at

CORRECTION: Thanks to Peter Lack—David Lack’s son, who was 16 at the time of that hike with his mother, father and 3 siblings—for these interesting details. I had previously reported here (and in Birkhead et al. 2014) that they were looking for and found the Brook Saxifrage.

In the Shadow of Men

BY: Bob Montgomerie, Queen’s University | 11 February 2019

In 1973, I was stranded for several days on a small island in Witless Bay off the southeast coast of Newfoundland. I had gone there several times already that summer, conducting seabird surveys for the Canadian Wildlife Service. Landing on Green Island was sometimes difficult and the days were short as the local cod fishermen—Bill White and Henry Yard—who took me out and back liked to do so on certain tides to make the landings less dangerous. As the season was getting on and I still had a large part of the island to census, I decided, one day in June, to stay overnight so that I could get in 3-4 times as many hours on the island than was possible on a single visit. I took a tiny pup tent, two days’ food and water, a small camp stove full of fuel, a sleeping bag and a change of clothes as I knew I’d get wet.

shadowMy best-laid plans were thwarted by a fierce, unexpected storm that came to shore that night and lashed the island for more than a week. The storm was so wild that it prevented both the fishermen and an RCMP helicopter from picking me up. Often I had to spend hours in my little tent to stay dry and to keep from being blown off the cliffs. To pass the time I slept, made plans to stretch out my meagre food supply, and organized my field notes. I also built nooses of fishing line to catch some murres in case I needed to eat a few to survive as the storm was showing no signs of letting up. When cooped up in my little tent, I read, several times, the only book I had taken with me, Jane Goodall’s In the Shadow of Man, which had just come out in paperback. That book—and the experience of being stranded and rescued—had a profound effect on me.

Three things about Goodall’s book were important to my development and outlook as a scientist. First, and foremost, this was the first book I had read by a woman biologist/naturalist [1], and it was just as good as all the others. I think that Goodall’s book more or less marked a turning point for biology that has transformed the role of women during the past 50 years. Prior to Goodall’s book, I had read many of the recent and now classic ‘popular’ books by and about naturalists—Tinbergen, Lorenz, Lack, Robert Ardry, George Schaller, Ernest Thompson Seton, Albert Hochbaum, Fred Bodsworth, James Fisher and Roger Tory Peterson, to name just a few—all by men.

At that time, I knew of excellent recent work by women ornithologists—MM Nice, Mary Willson, Mercedes Foster, Janet Kear, Susan Smith—published in the bird journals, but they were very much in the minority. I have been looking at the publications by female ornithologists in The Auk and The Condor over the last 135 years and the trend—and the exponential increase in female authorships since 1970—is shown on the graph below, reflecting a similar trend in The American Naturalist.


My own experience as an academic reflects this welcome pattern as well. My first group of four graduate students were all men, all of whom went on to academic positions at excellent universities. My last (in both senses of the word) four graduate students were all female. It’s too early to tell what career path they will take but one of them just got a tenure-track job. This change in the composition of my research group since 1980 does not reflect any conscious attempt on my part to train women scientists—everyone that I took on as a graduate student was simply the best applicant at the time. During my first decade teaching (1980s), most undergraduate biology students were male; when I looked out on my 48-student History and Philosophy of Biology class last week I could count only 9 men.

Stilt Sandpiper, Churchill 1972

Goodall’s book also reminded me how much fun it is to study animals close up, and how much better your insights can be when you can get extremely close to animals without seeming to disturb them. I enjoyed that aspect of studying seabirds that summer in Newfoundland, but also when studying both sandpipers and collared lemmings on the tundra at Churchill, Manitoba, the previous two summers. Such close observations of behaviours seemed to be important for testing hypotheses in the nascent field of behavioural ecology, especially where social interactions were concerned. Partly for that reason, I returned to the arctic with my newly-minted research group in 1980 as I knew the birds would be tame, could be watched at close distance, and could be followed for as long as we wanted on the open tundra. That was one of the reasons that we were able to document high levels of extrapair mating in Lapland Longspurs, years before DNA fingerprinting revealed that extrapair paternity was common in passerine birds [2].

Ike and Tina just outside my tent

Finally, I was amused that Goodall had named all of the chimpanzees that she watched. I knew that Lorenz and others had named their study animals but I always thought that that would not be acceptable in a serious scientific study. Goodall reminded me that there was nothing wrong with making research fun and entertaining. Right away I started to give names to the pairs of seabirds nesting near my tent—was I going a little stir crazy? For the local pairs of puffins, black guillemots, herring gulls and common murres, I chose the names of my favourite folk and rock couples—Ian and Sylvia, Ike and Tina, (Peter) Paul and Mary, Jim and Jean, and Chuck and Joni [3]. Years later, we often gave names to our favourite pairs of Lapland Longspurs, Snow Buntings and Rock Ptarmigan. And, in the early 1990s, when we studied Ruffs on Gotland in the Baltic, we named each of the males on every lek and used hand-drawn mug shots to identify them individually.

Ruffs on Gotland—copy of a page from my 1990 field notes on Gotland

Today (11 February 2019) is the UN-sponsored International Day of Women and Girls in Science, designed to celebrate and promote the roles of women in all of the sciences. While we have come a long way since Jane Goodall began working on chimpanzees, less than 30% of scientists worldwide are women, and there are still many barriers and sources of discrimination and gender bias in the sciences.

At the AOS meeting in Anchorage this year we will have some displays celebrating the roles of women in ornithology. For a long time, ornithology was largely a man’s game [4] but there have been some great, but relatively unknown, woman ornithologists in the past. I have tried to highlight some of their accomplishments on this blog [4]. In that same vein, I will devote all of March (Women’s History Month in the USA) to posts about the contributions of women to ornithology before Jane Goodall began studying chimpanzees.


  • Bronstein JL, Bolnick, DI (2018) “Her Joyous Enthusiasm for Her Life-Work…”: Early Women Authors in The American Naturalist. American Naturalist 192:655-663.

  • Burke T, Bruford MW (1987) DNA fingerprinting in birds. Nature 327:149–152.

  • Klopfer PH (1962) Behavioral aspects of ecology. New Jersey: Prentice-Hall, Inc.

  • Goodall JvL (1971) In the shadow of man. Boston: Houghton Mifflin.


  1. first book I had read by a woman biologist/naturalist: it’s only when writing this post today that I realized this. It certainly did not surprise me at the time.
  2. Chuck and Joni: my friends and I went to hear folk concert by the Mitchells at a coffee shop (either Penny Farthing or Riverboat) in Toronto’s Yorkville Village one night in 1967 or so. But the couple had broken up the day before and so a very nervous Joni did the gig on her own. She never looked back.
  3. extrapair paternity was common in passerine birds: see Burke and Buford (1987) for an early example
  4. largely a man’s game: see previous posts here, here, here, here, and here

IMAGES: all photos, the Ruff drawings, and the Auk/Condor graph by the author; American Naturalist graph modified from Figure 1 in Bronstein and Bolnick (2018)

CORRECTIONS: in the original post I forgot to add the Bronstein and Bolnick reference, the image sources, and the drawings of Ruffs. All added on 13 Feb 2019

Three French Hens

BY: Bob Montgomerie, Queen’s University | 24 December 2018

Tomorrow is Christmas Day and, like last year, I am spending the holidays in the north woods, a few km south of the southern tip of Algonquin Park, on the southern edge of the Canadian Shield. I wrote about the Twelve Days of Christmas song a year ago and am repeating that essay here with some new pictures and a few additions—including some details about those Three French Hens—that I have learned about over the past year.

I particularly like The Twelve Days of Christmas because the words are secular, even though there are myriad religious interpretations [1]. The song originated in an 18th century memory game, celebrating an annual period of drunkenness and merrymaking sandwiched between two religious feasts. Many of those twelve days are about birds that were prized for the table. In mediaeval England, this period following Christmas was presided over by the Lord of Misrule and in Scotland by the Abbott of Unreason, both titles that I would be proud to bear.


The words to this Christmas song were first published in English in the late 1700s as a rhyme in a book called Mirth without Mischief, likely derived from a much older French song of similar structure and content, Les Douze Mois. The now familiar tune was not written until 1905 by the English composer Frederic Austin who adapted it from a traditional English folk melody.

As you will recall—for by now it’s an ear worm that you can’t stop humming—the 12 days begin on Christmas Day with the partridge. On 5 or 6 of the following days, the gifts are birds, interrupted musically, thematically and enigmatically by those 5 golden rings. I have no idea why the first 7 gifts are birds, but I expect there are traditional and psychological reasons that have been claimed for this but they are probably all about food. There have also been many Christian interpretations of this song but really no evidence to support any of them. I find the secular interpretations to be far more interesting and valid.

In the almost 238 years since the rhyme was first published in English, there have been at least 20 different versions of the words, especially with respect to the birds. Some of these variants are undoubtedly Mondegreens [2], but they were often probably just attempts to make the words more relevant to a contemporary audience.

The PARTRIDGE—on the first day of Christmas— was always a partridge, except in Scott’s 1892 version where it was a “very pretty peacock.”  Some authors claim that the partridge was the Red-legged Partridge (Alectoris rufa) a very popular game bird that had just been successfully introduced to England from France in about 1770, and much more likely to perch in trees than the native and abundant Grey Partridge (Perdix perdix). But what about that pear tree, which again has been often claimed to have religious connotations. The French poem that may have been the basis for the English rhyme has a partridge representing the first month “Un’ Perdix Sole’. That version says that the bird flies in the woods (‘qui vol dans les bois’). The Perdix is the Grey Partridge, which in Old French was spelt ‘perdrix’ or ‘pertriz’, pronounced something very close to ‘pear tree’. I wonder if the English rhyme was originally ‘partridge and a perdrix’, though that would be two birds for day one. Nonetheless it seems to me quite likely that the pear tree was actually the perdrix, and had nothing at all to do with pears or trees.

On day 2, the TURTLE DOVES were French hens in one 1877 version, and the FRENCH HENS on day 3 were once ‘fat hens’ in 1864, and turtle doves in 1877. There’s a theme here as the first 3 birds were highly prized for the table, an excellent start to a period of feasting.

GallicRoosterBut why ‘French‘ hens? The Latin word for chicken is gallus and, as a result, the scientific name is Gallus gallus [3]. In Roman times, France was Gaul, and people who lived there were Gallic. It seems that the simple word association between the homonyms Gallus and Gallic irrevocably associated the fowl with France. Indeed, a rooster was often a decorative ornament on church bell towers in France during the Middle Ages, and the Gallic Rooster (see photo, right) was an important symbol during the French Revolution.

Bresse Gauloise

But also, when the Twelve Days rhyme was written, French hens were a prized table bird in both France and England. The breed Bresse Gauloise, for example, was sometimes called the ‘queen of poultry and the poultry of kings’. This breed originated in France in the late 16th century. La Fleche is also an ancient French breed from the Loire region of western France, and was renowned for its delicate flesh. During the 16th century hens from France were a luxury import from France. In the 19th century, the Houdan, another old breed from west of Paris, was one of the main meat breeds of France, and was imported to North America in 1865.

La Fleche

We humans are inordinately fond of eating chickens and a recent report suggests that the 60 billion chickens that we slaughter every year may turn out to be the paleontological signal of the Anthropocene. Of all the birds mentioned in the Twelve Days of Christmas, I doubt that anyone in 1800 could have predicted that the French hens and their kin would someday become the most abundant bird in the world, by at least an order of magnitude.

The CALLING BIRDS of day 4 are the most interesting to me as the original said ‘colly birds’ and subsequent variants said the birds were ‘canary’, ‘collie’, ‘colley’, ‘colour’d’, ‘curley’, ‘coloured’, ‘corley’, and finally ‘calling’ by Austin in 1909 published with his new tune. I am surprised no one ever suggested ‘collared’. The original ‘colly bird’ was the European Blackbird (Turdus merula) as ‘colly’ meant ‘black’ as in ‘coaly’, and is why border collies bear that name. The subsequent versions are undoubtedly the result of mis-hearings and misinterpretations.

The gift for day 5 in the original and modern version is GOLDEN RINGS but several sources claim that these are birds too, probably European Goldfinches, which were called goldspinks in the 1700s. Others have argued that these were Ring-necked Pheasants which have been claimed to have golden rings around their neck (but they don’t). The pheasant interpretation matches the culinary theme of the other 6 birds in the song, but the goldfinch was a popular cage bird in the 18th century. The melodic break in the song suggests a change of theme but the melody was added more than a century after the words.

The birds of days 6 and 7—the GEESE A-LAYING and the SWANS A-SWIMMING—round out the culinary theme before the song turns to dance providing some exercise after all that feasting, and chores that may have been neglected.

Here in the north woods the colly birds (and the only birds really calling) are Ravens, and the only ‘partridge’ is the Spruce Grouse, as all the geese, swans, doves, and goldfinches have departed for more southern winter quarters. The good news, this year, is that there are Pine and Evening Grosbeaks in the neighbourhood, as well as both species of redpoll. E-bird (map below) shows that I am well-situated (white star) to see Evening Grosbeaks in numbers, a bird I have seen only occasionally for the past 50 years.

Sightings of Evening Grosbeak Oct-Dec 2018 (from e-bird)

Counting the 5 golden rings, there are 28 individual birds in The Twelve Days of Christmas but I will be lucky to see even 28 individual birds on a day out in the winter woods here, where the temperature will be below freezing—and sometimes way below—for the next four months. That will not stop the 75 or more people who will gather in Algonquin Park for the Christmas Bird Count (CBC) on 29 December, where they will probably record fewer than 28 species [4] in a hard day’s work on foot, skis and snowshoes. This will be the 45th consecutive CBC for Algonquin Park and the 118th CBC since Frank Chapman started the count in 1900.

During the 19th century, the Christmas Side Hunt was a popular competition to gather game for the table during the 12 days of Christmas. Chapman, however, was a conservationist who saw great value in watching rather than hunting birds. That first CBC involved only 27 birdwatchers at 25 sites from Toronto, Ontario, to Pacific Grove, California, laying the foundations for what we now call citizen science.

French Hens (Houdans)


  • 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.


  1. myriad religious interpretations: see here, for example
  2. Mondegreen: Jimi Hendrix created a classic ‘Mondegreen’ when he sang (at least to my ears) “Scuse me while I kiss this guy” in his song ‘Purple Haze’, first released as a single in 1967. Rock lyrics are a rich source of Mondegreens—words or phrases that are misheard—as Sylvia Wright, who coined the term, did when she heard a Scottish ballad say “Lady Mondegreen” when it actually said “laid him on the green”.
  3. Gallus gallus: Linnaeus established this in 1758, but John Ray called them Gallus gallinaceus in 1676 and the name had clearly been in use for some time in England and Europe. Gallus gallus is, of course, the scientific name of the wild ancestor of the domestic hen, the Red Junglefowl of southeast Asia
  4. fewer than 28 species: that’s what I predicted for last year and they did indeed record that number, and 4704 individuals. Their sighting rate was 31 birds per party hour and that was well above the average of 25. That’s a lot of work (maybe 3 birds per hour per party), but a great day out.

Joe Grinnell’s Notes

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

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

Detail from a page in Linnaeus’s commonplace book

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

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

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

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


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

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

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

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

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

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

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

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


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

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

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

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


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

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

A Whiskey-Jack by Any Other Name

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

Canada Jay, Algonquin Park, 1968

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

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

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

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

from the AOU Checklist 1957

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

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

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

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

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

from the AOU checklist 1886

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

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


  • American Ornithologists’ Union (1886) The code of nomenclature and check-list of North American birds. New York: American Ornithologists’ Union.
  • American Ornithologists’ Union (1957) Check-list of North American birds. Ithaca, N.Y.: American Ornithologists’ Union.
  • Brisson M-J, Martinet FN (1760) Ornithologie, ou, Méthode contenant la division des oiseaux en ordres, sections, genres, especes & leurs variétés (t.2 ). Parisiis: Ad Ripam Augustinorum, apud Cl. Joannem-Baptistam Bauche, bibliopolam, ad Insigne S. Genovesae, & S. Joannis in Deserto.
  • Hearne S (1795) A journey from Prince of Wales’s Fort in Hudson’s Bay to the Northern Ocean. London: Strahan and Cadell.
  • Houston CS, Ball T, Houston M (2003) Eighteenth-century Naturalists of Hudson Bay. Montreal: McGill-Queen’s University Press.
  • Linné CV (1766) Caroli a Linné. Systema naturae : per regna tria natura, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis / (t.1, pt. 1 (Regnum animale)), 12th edition. Holmiae :Impensis direct. Laurentii Salvii.
  • Pennant T (1784-85) Arctic Zoology, 2 vols. London: Henry Hughs
  • Rutter, R.J. 1969. A contribution to the biology of the Gray jay (Perisoreus canadensis). Canadian Field-Naturalist 83: 300-316.
  • Strickland D (1969) Écologie, comportement social et nidification du Geai Gris (Perisoreus canadensis). Master’s Thesis, Univ. Montréal, Montréal.
  • Strickland D (2017) How the Canada Jay lost its name and why it matters. Ontario Birds 35: 2-16.
  • Swainson W, Richardson J (1831) Fauna boreali-americana, or, The zoology of the northern parts of British America: containing descriptions of the objects of natural history collected on the late northern land expeditions, under command of Captain Sir John Franklin, R.N. Part second, the birds. London: John Murray.


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

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

Professor Bumpus and his Sparrows

Guest Post

BY: Ted R. Anderson | 5 March 2018

Possibly the most influential ornithological paper published inNorth America in the 19th century was actually written by an invertebrate embryologist who was not even a member of the American Ornithologists’ Union.  The paper “The elimination of the unfit as illustrated by the introduced sparrow, Passer domesticus” was written by Professor Hermon Carey Bumpus at Brown University in Providence, Rhode Island  It was actually the second of two interesting papers that Bumpus published on the recently introduced house sparrow, but more on these below.

Hermon Bumpus

Bumpus was born in Maine in 1862, and entered Brown in 1879 to study biology, graduating in 1884.  In 1886, he accepted a professorship at Olivet College in Michigan, a position he left in 1889 to complete a doctorate at the newly established Clark University, where he received the first PhD awarded by that university.  In 1890 he returned to Brown as assistant professor of zoology and was promoted to professor of comparative anatomy two years later.  He left Brown in 1900 to become assistant to Morris Jessup, president of the board of the American Museum of Natural History in New York City.  A year later Jessup promoted him to become the museum’s first director.

In 1911 Bumpus moved into academic administration as business manager of the University of Wisconsin, a position he held until 1914.  He then moved to Tufts College (now University) as President from 1915 to 1919.  He resigned from Tufts to pursue his interest in building or remodeling homes including a Philippine bungalow on Long Island Sound (constructed of Philippine lumber from the Philippine Hall at the St. Louis Exposition), an Italian villa in a Boston suburb and the King Caesar House in Duxbury, Mass.  Bumpus died in Pasadena, California in 1943.

pexels-photo-460960.jpegWhile teaching at Brown, Bumpus spent his summers conducting research on the development of marine invertebrates at the Marine Biological Laboratory at Woods Hole, Massachusetts, where he also served as assistant director from 1893 to 1895, and as director of the Biological Laboratory of the US Fish Commission.  In 1896 he presented the inaugural lecture in a summer seminar series at the Marine Biological Laboratory entitled “The variations and mutations of the introduced sparrow, Passer domesticus”, which was published in 1898.

In that 1898 paper he was undoubtedly the first scientist to suggest that the introduction of house sparrows and their subsequent rapid spread across North America represented a huge experiment that could be used to study Darwinian natural selection.  Taking advantage of that natural experiment, he compared the size, shape and coloration of 868 sparrow eggs from Massachusetts with an equal number of sparrow eggs from England, to test the hypothesis that the rapid population growth of sparrows in North America would result in relaxed selection.  Without the benefit of statistical analysis—Francis Galton and Karl Pearson were just then developing some rudimentary statistical tests—he concluded from his graphs that eggs from Massachusetts were shorter and more variable in size and coloration than eggs from England.  He also raised the question of whether the observed differences were phenotypic (‘ontogenetic’) or adaptive (‘phylogenetic’) and suggested that a common garden experiment would be needed to differentiate between these alternatives.

Bumpus’s graph of the length of house sparrow eggs from North America (dotted line) and Europe (solid line) [1]

On 1 February 1898, a winter storm in Providence provided Bumpus with the material for another summer lecture at Woods Hole, which he then published.  After the storm, 136 immobilized sparrows were brought to Bumpus’s anatomy lab, where 72 subsequently revived but the remaining 64 died.  Bumpus identified the sex and measured nine morphological traits of each bird.  Bumpus concluded from his graphs that males survived better than females and that shorter, lighter birds with longer legs, wings and sternums and larger brain size (“skull width”) also survived better. He concluded that his analyses showed:

Natural selection is most destructive of those birds which have departed most from the ideal type, and its activity raises the general standard of excellence by favoring those birds which approach the structural ideal.

…the birds which perished have certain average structural peculiarities which distinguish them from the survivors, and that the intensity of selective elimination has been felt most by birds of extreme structure [2]

In his 1899 publication, the entire dataset is reproduced in an appendix, thereby permitting many other evolutionary biologists, as well as innumerable students in evolution classes, to analyze Bumpus’s data statistically.  Harris published the first  statistical analysis, and at least ten other papers have been published since then, including papers by John Calhoun, Peter Grant , Richard F. Johnston and colleagues, and one by Russell Lande and Steven Arnold.  Increasing complex and sophisticated statistical analyses were employed in these papers, and the conclusions of the various authors differ from those of Bumpus and from each other, in part due to the fact that many of the analyses use only subsets of the original data.

I do not know of another dataset of birds that has been subjected to so many analyses and so many different interpretations The history of reanalysis of Bumpus’s data is a nice example of a century of progress in both statistics and evolutionary biology


  • Anderson TR (2006) Biology of the Ubiquitous House Sparrow, from Genes to Populations. Oxford: Oxford University Press
  • Bumpus HC (1898) The variations and mutations of the introduced sparrow, Passer domesticus. Biological Lectures Delivered at the Marine Biological Laboratory of Woods Holl, 1896-1897, pp. 1-15.
  • Bumpus HC (1899) The elimination of the unfit as illustrated by the introduced sparrow, Passer domesticus. Biological Lectures from the Marine Biological Laboratory of Woods Holl, Mass. 1898, pp 209-228.
  • Calhoun JB (1947) The role of temperature and natural selection in relation to the variations in size of the English sparrow in the United States. American Naturalist 81:203-228.
  • Grant PR (1972) Centripetal selection and the house sparrow. Systematic Zoology 21:23-30.
  • Harris JA (1911) A neglected paper on natural selection in the English sparrow. American Naturalist 45:314-319.
  • Johnston RF, Niles DM, Rohwer SA (1972) Hermon Bumpus and natural selection in the house sparrow Passer domesticus. Evolution 26:20-31.
  • Lande R, Arnold SJ (1983) The measurement of selection on correlated characters. Evolution 37:1210-1226.


  1. graph: Bumpus 1898 page 5
  2. quotation: Bumpus 1899 pages 217 and 218

Birthday wishes

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

DarwinsgraveMr Charles Darwin
Westminster Abbey
20 Deans Yd
London SW1P 3PA

My Dear Charles (if I may)

Happy 209th birthday! 

I know that a few people have written to you [1] in the 132 years since you shuffled off this mortal coil, but I thought it high time we brought you up to date on the research inspired by those drab little finches that you collected in the Galápagos Islands. I am not really sure how long it takes for letters to reach you at your new address [2] but the current evidence suggests that it might take forever. But still…

You presumably know about the several Galápagos expeditions that took place during your lifetime. As soon as John Gould had figured out that there were actually 12 species of finches in your collection, and that there were different combinations of species on different islands, the floodgates were opened.

You will remember poor old Thomas Edmonstone [3] who went to the islands on HMS Herald in 1845. You had asked him specifically to collect finches from as many different islands as he could visit and to keep careful records as to which specimens were from which island, as you had not properly labelled all of your own finch specimens. Sadly, he died on that voyage and his records were virtually indecipherable.

Then Simeon Habel went out in 1868 and stayed for 6 months, making an extensive bird collection that ended up in Vienna. Percy Sclater and Osbert Salvin [4] published a little about that collection. Finally, your old nemesis, Luis Agassiz [5], went to the islands for only 9 days in 1873 but his crew still collected a lot of specimens for the Harvard Museum of Comparative Zoology. Agassiz, as you no doubt expected, could not see what all the fuss was about over the finches and remained a devout creationist.

It has always been pretty clear to me that the finches fascinated you. Even in your notes from the Beagle voyage you could already tell that they were closely related to some mainland species and realized that they were all related, even if, at the time, you thought that many of the finches on different islands were just morphologically differentiated races of the same species. As you said in your ‘Ornithological Notes‘:

These birds are closely allied in appearance to the Thenca of Chile or Callandra of la Plata. In their habits I cannot point out a single difference; — They are lively inquisitive, active run fast, frequent houses to pick the meat of the Tortoise, which is hung up, — sing tolerably well; are said to build a simple open nest. — are very tame, a character in common with the other birds…I have specimens from four of the larger Islands; the two above enumerated, and (female. Albermarle Isd.) & (male: James Isd). — The specimens from Chatham & Albermarle Isd appear to be the same; but the other two are different. In each Isld. each kind is exclusively found: habits of all are indistinguishable… When I see these Islands in sight of each other, & possessed of but a scanty stock of animals, tenanted by these birds, but slightly differing in structure & filling the same place in Nature, I must suspect they are only varieties. (Darwin 1963)

In 1888 and 1891, Zera L. Tanner took several naturalists (including Aggasiz’ son Alexander) on his fisheries research ship USFC Albatross to the islands to collect specimens. The bird specimens in those collections, as well those of Habel and Bauer and Adams in 1891, were studied by the great American ornithologist Robert Ridgway [6] and published in his Birds of the Galapagos Archipelago in 1896. Ridgway recognized that despite all of the previous visits to the islands, not much was really known about the finches still:

Not a single island of the group can be said to have been exhaustively explored, and few of the species are known in all their various phases in fact, many are known only from a few specimens in female or immature dress. No observations have been made upon the attitude the different species of Geospiza maintain toward one another tending to show how far the differences observable, or thought to be observable, in dried specimens indicate the actual grouping in species of living individuals. The anomaly of individuals adult as to plumage but with bills suggesting immaturity’, and of others which show exactly the reverse, remains to be explained and there are other questions which only protracted field-studies by a competent investigator can decide. Until all these present mysteries are solved, theories and generalizations are necessarily futile. (Ridgway 1897: 459-460)

Rollo Beck

The indefatigable Rollo Beck of California was the next ornithologist to make collections in the Galeapagos when he was sent there by Lord Walter Rothschild [7]  specifically to collect tortoises that were sent back to Rothschild’s collection at Tring (UK). Beck was sent back to the islands in 1905 to collect for the California Academy of Sciences, and stayed for a whole year. He shipped 78,000 specimens (including 8688 birds [8]) back to the Cal Academy, a third of which were finches.

Percy Lowe went out in 1936 but declared in his paper published in The Ibis that the finches were all a kind of hybrid swarm with no clear evidence of speciation. He did, however, call the birds ‘Darwin’s Finches’ and you will undoubtedly be delighted to know that that name endures to this day despite a few subsequent publications calling them ‘Galapagos Finches’.

LackDFIn 1938-39, a school teacher named David Lack spent four months on the islands studying the behaviour of the fiches, followed by a few months studying the extensive collection of specimens at the Cal Academy. Lack was already a well-known, if amateur, ornithologist having participated in some ornithological expeditions and published a few papers. Lack was appointed Director of the Edward Grey Institute of Field Ornithology in 1945, a position he held until he died in 1973. In 1940 he published the results of his Galápagos studies in a paper in Nature and later, in 1947, in a book called Darwin’s Finches. Lack really established the finches as model organisms for the study of evolution.

To some extent inspired by Lack’s book, Robert Bowman [9]—a Queen’s University graduate—went first went to the Galápagos in 1956 and returned several times to conduct a series of studies designed in part to show that Lack was mistaken about the role of competition in the evolution of the finches. Bob did his PhD at UC Berkeley, carefully describing the skulls and jaw musculature of the finches. Later he studied their songs and concluded that those species that coexisted in the same island differed as much in their songs as they did in their bills, and that the songs probably reduced the incidence of ‘cross-breeding’ as you called it.

Finally (though not actually the end of the story), a personal note. In the fall of 1973, my PhD advisor, Peter Grant, who was then studying competition in microtine rodents, came into our grad student office and asked me if I’d like to go to the Galápagos to suss out the finches as a possible focus for my thesis research. I had just started a PhD program at McGill University a couple of months earlier and was thinking then that I wanted to return to Newfoundland or the high arctic, as I had worked studying seabirds for the Canadian Wildlife Service in both places in the preceding summer. Peter and his postdoc BeakIan Abbott were going to the Galápagos for a few weeks mainly to study the interactions between the birds and the seeds of Tribulus, which formed a large part of their diet. For several reasons I declined, but Peter (with Ian and his wife, Lynette) went and was enchanted, thereby beginning the longest running—and today probably best known—series of studies of ‘your’ finches. His work was the subject of a Pulitzer prize winning book called The Beak of the Finch. I am certain that you would be stunned by the quality and breadth of the work that Peter, his wife Rosemary, and their students and colleagues have accomplished in the intervening 45 years.

I am actually in Ecuador today in the cloud forest nw of Quito, watching birds with a couple of old friends, Tim Birkhead (sperm competition [10], history of ornithology) and David McDonald (manakins). Tim and I will fly [11] to Baltra on Thursday the 15th to tour the islands for a few days. I will send you some photos and field notes next Monday from our trip. That post might be delayed by a day or two depending upon the quality of the internet connection [12] on our boat or in Guayaquil. I know you did not get to the mainland of Ecuador in 1835 but I think that history has shown that your time was better spent in the Galápagos.

Yr obd svt

Bob Montgomerie


  • Darwin CR (1963) Ornithological notes. Barlow N. , editor. British Museum (Natural History) Bulletin, Historical Series 2: 201–278.
  • Lack D (1947) Darwin’s Finches. Cambridge: Cambridge University Press.

  • Lowe PR(1936) XV.—The finches of the Galapagos in relation to Darwin’s conception of species. Ibis 78: 310–321, doi:10.1111/j.1474-919x.1936.tb03376.x

  • Ridgway R (1897) Birds of the Galápagos archipelago. Proceedings of the U. S. Nayional Museum 19:459–670.

  • Sclater PL, Salvin O (1870) Characters of new species of birds collected by Dr Habel in the Galapagos Islands. Proceedings of the Zoological Society of London 38:322–323.

  • Ridgway R (1897) Birds of the Galápagos archipelago. Proceedings of the U. S. National Museum 19:459–670.



1. see photo at the top, but also, if you modern readers of this post are familiar with the what3words app or website, Darwin is at, or very close to, salsa.snap.finger

2. see for example a letter from Jerry Coyne on the Oxford Press blog in 2009 here; and a letter from Frank Gannon in EMBO Reports, also in 2009, here

3. 1825-1846; sometimes spelled Edmonston; he was accidentally shot in the head after landing in Peru, right after visiting the Galapagos 

4. Sclater (1829-1913) and Salvin (1835-1898) were both prominent 19th century English ornithologists who were original members of the BOU and editors of The Ibis. Together they published the first paper to appear in The Ibis

5. Agassiz(1807-1873) was born and raised in Switzerland but moved to the USA in 1847 and eventually became professor of zoology at Harvard where he founded the Museum of Comparative Zoology.

6. Ridgway (1850-1929) was one of the founders of the AOU (now AOS) and was president of the AOU from 1898-1900

7.  Rothschild (1868-1937) was a wealthy collector who donated his Tring estate to the British Museum, where it is now a division of the Natural History Museum and sold his bird specimens to the American Museum of Natural History, thereby establishing the AMNH as a leading ornithological centre worldwide.

8. About 2500 of these were finches in the genus Geospiza alone (data from VertNet accessed on 4 Feb 2018)

9. Bowman (1925-2006) first went to the islands as a grad student, then several more times during his long career as a professor at San Francisco State University until he retired in 1988.

10. It wasn’t until 1970 that Geoff Parker added this interesting way that males compete to your list of important mechanisms of selection.

11. I don’t know if you are aware that we humans can now fly from place to place in giant buses with wings called airplanes, making it possible to get from my home in Canada to the Galápagos in less than a day. The first airplane flew to the Galapagos in 1934 to rescue a naturalist who had contracted appendicitis.

12. I don’t even know where to start explaining this. It’s like the mail without the paper, and messages are transferred instantly (which is both a boon and a curse) across the ether, so you are actually likely to encounter them wherever you are.

Uncle Bill’s Eggs

BY: Bob Montgomerie, Queen’s University | 13 Nov 2017

Yesterday (12 November) marked the anniversary of the discovery, in 1912, of the remains of Robert Falcon Scott’s ill-fated Terra Nova Expedition to the South Pole. The story of that expedition’s side-trip to collect Emperor Penguin eggs is well-known, celebrated in myriad books, articles, documentary films and exhibitions. As is often the case with scientific exploration and discovery, there is a less-well-known backstory that raises some interesting questions about the more celebrated account.

Edward Adrian Wilson was the doctor, zoologist, and artist on Scott’s two famous Antarctic expeditions—Discovery in 1901-04 and Terra Nova in 1910-12. Wilson was a well-connected English artist and ornithologist, elected as a member of the BOU in 1900. Little surprise, then, that the prominent English ornithologist Percy Sclater [1] suggested to Wilson that he apply to be Junior Surgeon and Zoologist on Scott’s Discovery Expedition. On returning from that first expedition, Wilson worked as Field Observer for the Grouse Disease Commission in Scotland, but that’s a story for another day.

Wilson’s painting of Emperor Penguins with chicks at Cape Crozier in September 1903

On the Discovery expedition, Wilson visited the Emperor Penguin colony at Cape Crozier where he made an extensive study of their breeding biology. In his published report of 1907, he mentions that:

The possibility that we have in the Emperor Penguin the nearest approach to a primitive form not only of a penguin but of a bird, makes the future working out of its embryology a matter of the greatest importance. It was a great disappointment to us that although we discovered their breeding ground. and although we were able to bring home a number of deserted eggs and chicks, we were not able to procure a series of early embryos by which alone the points of particular interest can be worked out…The whole work [of getting eggs for embryological study] no doubt would be full of difficulty, and it is with a view to whom the opportunity may occur in the future, that this outline has been added of the difficulties that would surely beset their path. [2]

Having provided all of the details needed to procure those precious embryos, Wilson was determined to get them during his second expedition to the Antarctica with Scott. But why were those embryos so important?

At the turn of the twentieth century, Ernst Haeckel’s Biogenic Law that ‘ontogeny recapitulates phylogeny’ was still very current. Wilson thus thought that an examination of the embryonic development of a bird thought to be very ‘primitive’ [3]—the Emperor Penguin—might shed some light on the origin of birds. Thus Wilson felt that the study of Emperor Penguin embryology might reveal some of the details about how birds evolved from reptiles, as had earlier been suggested by Sir Thomas Huxley and others. In the same volume as Wilson’s report from the Discovery expedition, William Plane Pycraft [4] wrote about penguin anatomy, based on specimens from the Discovery expedition. Pycraft laid the foundation for Wilson’s quest for the embryos when he speculated that:

All that can be gleaned from fossils, then, is that penguins have probably descended from birds which possessed full powers of flight, and this probability becomes converted into a certainty when the embryological evidence comes to be examined. But the question of the precise affinities of this group must still be regarded as an unsolved problem, the intense specialisation which these birds have undergone obliterating much of the necessary evidence. [5]

Wilson may also have thought that those embryos might answer the question about the origins of flightlessness on the penguins—had they evolved from flying birds, for example—by comparing their embryological development with that of other ‘primitive’ non-passerine birds like ducks.

In an 1887 paper, Mikhail Menzbier had also speculated that the extant birds might have evolved from two independent lineages leading from the reptiles, one to the flightless penguins and the other to the flying birds. We don’t know if Wilson even knew about that idea but a detailed study of the penguin’s embryology might have helped to resolve that issue as well.

Wilson (left), Bowers (middle) and Cherry-Garrard (right) in August 1911, shortly after returning from their trek to collect the eggs

During the Terra Nova expedition, Wilson’s fellow explorers found him to be very companionable and someone they could confide in—they called him ‘Uncle Bill’. Near the start of that expedition, in July 1911, Wilson, Henry Robertson “Birdie” Bowers [6] and Apsley Cherry-Garrard and made a horrendous winter trek—95 km each way, in the dark with extreme cold—from Cape Evans to the Cape Crozier colony where they obtained five incubated eggs that Wilson thought could be used for embryological study. Though Wilson and Bowers died with Scott and two others, Cherry-Garrard returned three of those eggs [7] to England where they were later dissected by three different embryologists over the next 20 years. One of those embryologists concluded—contrary to Wilson’s hopes—that those eggs: have not contributed much to the understanding of the embryology of penguins.

“Uncle Bill” Wilson’s Emperor Penguin eggs from Cape Crozier 1911

Strangely enough, the question that Wilson hoped to answer by obtaining those embryos had already been answered by the dissection of Gentoo and Adélie Penguin eggs collected by Robert Neal Rudmose-Brown [8] and James Hunter Harvey Pirie [9] on the Scottish National Antarctic Expedition of 1902-04 . The embryos in those eggs were studied by anatomists David Waterston and Auckland Campbell Geddes at Edinburgh University who concluded, in October 1909, that:

With regard to these developmental facts the question arises:— Is the duck’s or the penguin’s wing the more direct descendant of the common ancestor; or have they both diverged from the common stock approximately equally, but in opposite directions?

Embryology alone cannot answer this question, but the evidence is clear in this, that the fore limb of the penguin in its development goes through a progressive and continuous series of stages along one unbroken line…So that the answer to our question, so far as the embryological evidence is concerned, must be that the wings of both these birds are different from the ancestral wing, and that the differentiation has been in opposite directions and that the common ancestor was a flying bird of a somewhat primitive type depending in large measure for the spread of its wing upon bone and muscle. [10]

How did Wilson not know about this work, published in the Transactions of the Royal Society of Edinburgh, more than 8 months before the Terra Nova expedition began? Quite possibly he was too busy with his grouse research and illustration commissions to follow the recent publications, though that seems highly unusual given the intense interest in Antarctic exploration in general and penguins in particular in the early 1900s. Possibly, even if he knew about it, he might have felt that the work on Adélie and Gentoo Penguins could not actually answer the question because he may have felt that those species were not primitive enough. The Emperor is the only penguin that breeds in the Antarctic winter, and there was a notion, in those days, that this indicated that it was the most primitive bird.

Whatever the motives behind Wilson’s quest for Emperor Penguin eggs, his studies of their breeding biology is an outstanding early example of research into the breeding cycle, parental care, and offspring development of any bird.


  • Cherry-Garrard A (1922) The Worst Journey in the World. London: Chatto and Windus
  • Haeckel E (1866) Generelle morphologie der organismen [General Morphology of the Organisms]. Berlin: G. Reimer. (Accessed December 3, 2013).
  • Menzbier M (1887) Vergleichende osteologie der pinguine in anwendung zur haupteintheilung der vogel. Bulletin de la Société impériale des naturalistes de Moscou 1: 483-587
  • Mossman RC, Pirie JHH, Rudmose-Brown RN (1906) The voyage of the Scotia, being a record of a voyage of exploration in the Antarctic Seas. London: C. Hurst
  • Peaker M (2014) In Search of a Penguin’s Egg. Why? Zoology Jottings blog posts on 8 April and 6 June 2014. retrieved online on 12 Nov 2017 at and
  • Pycraft WP (1907) On some points in the anatomy of the Emperor and Adélie penguins. Section III pp 1-28 in Bell FJ, Fletcher L (eds) National Antarctic Expedition 1901-1904. Natural History Volume II. Zoology (Vertbrata : Mollusca : Crustacea). London: British Museum (Natural History) [available here]
  • Waterston D, Geddes A (1910). X.—Report upon the Anatomy and Embryology of the Penguins collected by the Scottish National Antarctic Expedition, comprising: (1) Some Features in the Anatomy of the Penguin; (2) The Embryology of the Penguin: A Study in Embryonic Regression and Progression. Transactions of the Royal Society of Edinburgh 47: 223-244
  • Seaver, G (1933) Edward Wilson of the Antarctic. Naturalist and Friend. London: John Murray
  • Wilson EA (1907) Aves. Section II pp 1-121 in Bell FJ, Fletcher L (eds) National Antarctic Expedition 1901-1904. Natural History Volume II. Zoology  (Vertbrata : Mollusca : Crustacea). London: British Museum (Natural History) [available here]
  • Wilson EA (editor) (1908) National Antarctic Expedition 1901-1904. Album of Photographs and Sketches. London: Royal Society


  1. Sclater was of the founders of the BOU and the first editor of The Ibis
  2. quotation from Wilson 1907: 31
  3. we try not to use that term ‘primitive’ anymore when talking about species because it is really traits that might be ‘primitive’ (i.e, present in a common ancestor) or derived when comparing two species
  4. in 1907 Pycraft was on the staff of the British Museum (Natural History)
  5. quotation from Pycraft (1907)
  6. he was called “Birdie” by his fellow expeditioners not because he had any interest in birds, but because he looked a bit like a bird with his red hair and beak of a nose
  7. two of the eggs broke when the trekkers climbed a cliff to begin their journey back to Cape Crozier
  8. Rudmose-Brown wasa botanist who was appointed lecturer in geography at the University of Sheffield in 1907
  9. Pirie was a bacteriologist and medical doctor
  10. quotation from Waterston and Geddes (1910); later reproduced as pp 37-58  in Volume IV of Report on the scientific results of the voyage of S.Y. “Scotia” during the years 1902, 1903 and 1904, under the leadership of William S. Bruce, published in 1915 [available here]

IAMGES: all images are in the public domain, available from Wikimedia Commons

Tools for Studying Birds

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

I bought a new pair of binos last week, from the incomparable Pelee Wings Nature Store near Point Pelee (the subject of a recent blog post) in Ontario. This is my 7th pair in more than half a century of watching and studying birds, and maybe the best (Swarovski Pocket CL 8×25); certainly the finest for their small size.

Binoculars are such an important tool for bird study that you could not really be a field ornithologist today without them. For too long, I relied on cheap bins until my friend (and at the time, postdoc), Geoff Hill, admonished me for using a toy to do professional work. And he was right—the Bushnell Elites (ca 1993) that he shamed me into buying allowed me to read color bands and examine individual plumage variation like never before.

Binoculars were not invented by or for birders, but eventually became the quintessential, discipline-defining tool for ornithologists. In his 1997 book, Image and Logic, the experimental physicist and science historian, Peter Galison, suggested that tools might be the main engine of scientific revolution, and not ideas as had earlier been suggested by Thomas Kuhn (1962) in his classic book The Structure of Scientific Revolutions.

Binoculars from the American Civil War

People studying birds slowly added binoculars to their field kits when they became commercially available but there is no indication that binoculars were in any way revolutionary for ornithology. Binoculars (as opposed to binocular telescopes) came on the market in the mid-1800s and at first figured prominently in military and astronomical applications, but not in bird studies. Even Edmund Selous’s 1900 classic Bird Watching, which arguably invented the hobby,  makes no mention of binoculars and the first reference I can find for their use in a bird study does not appear until 1923.

What other iconic bird study tools might have spawned revolutions in ornithology? Certainly many great discoveries about birds have been made with light and electron microscopes, tape recorders, computers and software, and DNA sequencers, but none of these were invented for or used almost exclusively by ornithologists.

Here is my short-list for essential tools that ornithology ‘owns’ in addition to binoculars and spotting scopes—tools that I think revolutionised ornithology:

  • metal and colour bands (rings): numbered metal bands were first made and used by a Danish schoolteacher, Hans Christian Mortensen, in 1899; colored markers (silver threads) were used by Audubon in 1803 but color bands as we know them today appear to have been first used in 1909 when Louis Gain (1913) put “some celluloid rings of various colors” on the legs of Adelie Penguins on Petermann Island, Antarctica; the rest, as they say, is history.
  • mist nets: mistnets were in widespread use to catch birds for food in Japan for at least three centuries before Oliver L. Austin used them to catch migrants in 1947. (Was he the first ornithologist to use these nets to study birds?) By the 1960s mist nets were in widespread use at banding (ringing) stations in North America and Europe, and had become an essential tool for field ornithology.
  • sound spectrographs, sonographs, sonograms: although developed at Bell Labs during WWII to break codes and identify aircraft by their sounds, even the first paper reporting on the technology showed spectrograms of five bird species (Potter 1945). By 1948, bird researchers from all over the world were ordering Sona-graphs from Kay Electric Co. (an offshoot of Bell labs).
  • radio transmitters and telemetry/geolocators/PIT and RFID tags/MOTUS: there are myriad electronic devices that can be attached to birds to find out where they are or have been. First used in the early 1970s for bird studies, these devices have been instrumental in determining both local movements and long distance migrations.
  • parabola/shotgun microphones: while the principle of focusing sound/light/radiation with a parabola was know for centuries, the first parabolas for bird song recording were made at Cornell University in the 1930s. In the 1960s, Dan Gibson, a wildlife cinematographer from Toronto, marketed a plexiglas version that became an essential tool for recording bird songs.
  • DNA fingerprinting (multilocus, microsatellite): first developed by Alec Jeffreys in 1984, and applied immediately to a human immigration case involving disputed family membership (Jeffreys 1985). It took only a couple of years before the first paper was published using DNA fingerprints to evaluate paternity in a wild bird, the House Sparrow (Wetton et al 1987). Paternity analysis certainly revolutionised studies of bird mating systems and mate choice.
  • portable color spectrometers (spectroradiometers): while ornithologists are a tiny fraction of the scientists who use relatively inexpensive, portable spectroradiometers in their research, their introduction in the early 1990s revolutionised the study of bird coloration
  • ebird: this online checklist system, launched in 2002 by the Cornell Lab of Ornithology and the National Audubon Society is already revolutionising our view of bird distribution and migration patterns in the western hemisphere (example here)

We will explore the history of these tools in more depth later. I am sure there are other tools that have changed the history of bird study, and I welcome your suggestions. With respect to ornithology, we know that both Kuhn and Galison were right (see also Dyson 2012), as the revolutions that have shaped the discipline have been fuelled by both new ideas and new tools.


Dyson FJ (2012) Is science mostly driven by ideas or by tools? Science 338:1426

Gain L (1913) The penguins of the Antarctic regions. Smithsonian Institution Annual Report 1912:473-482

Galison P (1997) Image and Logic: A material culture of microphysics. University of Chicago Press, Chicago.

Jeffreys AJ (1985) Positive identification of an immigration test-case using human DNA fingerprints. Nature 317:818-819

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

Potter RK (1945) Visible patterns of sound. Science 102:463-470

Selous E (1901) Bird Watching. JM Dent & Company, London.

Wetton JH, Carter RE, Parkin DT, Walters D (1987) Demographic study of a wild house sparrow population by DNA fingerprinting. Nature 327:147-149

IMAGE: binoculars from