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.

Joe Grinnell’s Notes

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

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

Detail from a page in Linnaeus’s commonplace book

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

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

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

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


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

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

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

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

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

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

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

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


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

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

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

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


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

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

A great store of fowle

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

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

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

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

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

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

Clets on St Kilda

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

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

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

Digges Island 1952 photo by Les Tuck

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

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

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

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


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


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

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

Why Woodpeckers are Scarce in the North

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

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

Wallace in 1862

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


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

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

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

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

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

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

Great Spotted Woodpecker

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


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

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

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

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


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

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

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

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

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


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

Serendipity 101

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

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

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

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

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

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

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

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

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

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

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

Ernst Mayr (right) in New Guinea 1928

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

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


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

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


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

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

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

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

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

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

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

IMAGES: all from Wikipedia, in the public domain