A Vacation Like No Other

[Note to readers: posting here has been on hiatus for the past couple of months as I have been in the field but also because this site will shut down in the next month or so, as the AOS launches a new website (here). Most, or all, of the current ‘history’ pages on this site will eventually migrate to the History Committee page on the new AOS site. This will be the final blog post in the current series that began here more than two years ago. In future I will contribute occasional posts to the omnibus AOS blog called Wing Beat. Thanks for reading and commenting on the 100 or so posts that have appeared on this site.]

BY: Bob Montgomerie, Queen’s University | 27 September 2019

Back in the 1980s, one of my graduate students and I split the cost of an Ontario Lottery ticket. We knew that the chances of winning were vanishingly small (p<0.000001) but the jackpot had risen to $24 million—real money in those days. It was a cheap and fun investment that could really help my research program if we won. We spent the next couple of weeks in the field, often chatting about what we might do with the winnings.

We eventually decided that, if we won, we would buy a nice plot of land in the arctic or the tropics, and build a field station. We figured there would be enough money to allow us to employ and fund a dozen or so field biologists to work at the station—people who would be engaging, creative and productive, with the freedom to explore whatever interested them. We did not win that particular lottery but both of us realized our dream by supervising some outstanding graduate students and postdocs in the intervening 30+ years, and working with them in the field.

E. H. Harriman

In 1899, Edward Harriman was faced with an opportunity like our dream of winning that lottery, but he already had the funds to make it happen. Harriman was what we used to call a railroad tycoon, and was one of the richest and most powerful businessmen in the United States. By 1899, when he was 51, Harriman was director of the Union Pacific Railroad and already fabulously wealthy. He achieved that success through incredibly hard work, combined with a keen mind and outstanding business acumen. The hard work, though, had taken its toll on his health, so, in January 1899, his doctor recommended a long vacation.

For Harriman, though, the idea of lying on a beach, visiting European cathedrals, or golfing for a month or two held little interest. But he knew he had to get away from the day-to-day running of his businesses for a while. His first thought was to spend a few weeks hunting Kodiak bears in Alaska, something he had always wanted to do. As often happens with type ‘A’ personalities, though, that simple ‘idea’ soon blossomed into a gargantuan endeavour involving huge expenses, dozens of people, and complex travel arrangements. Just as we had done 80 years later, Harriman thought it would be incredible to gather together some of the best field scientists in America and take them to an interesting place, all expenses paid and no expense spared.

To assemble a coterie of top scientists—geologists, historians, anthropologists, zoologists and botanists—Harriman, in March 1899, asked his friend C. Hart Merriam at the U.S. Department of Agriculture for help. Merriam was a founding member of the AOU (now AOS), the brother of Florence Merriam Bailey (see post here), and one of the top American zoologists of the day. Within three weeks, Merriam had convinced an incredible array of participants to join the Harriman Expedition, due to depart Seattle at the beginning of June. In addition to Merriam, these included ornithologists Robert Ridgway, Louis Agassiz Fuertes, Charles Keeler, Leon J. Cole, and Albert K. Fisher, as well as 10 other biologists, 5 geologists/geographers, and a dozen artists and photographers among others. The naturalists/authors John Burroughs, John Muir and George Bird Grinnell added to the sterling cast of explorers. Including his family members, a chaplain, taxidermists, cooks and the ship’s crew, a total of 128 people participated in the expedition.

Some of the expedition members at the abandoned Cape Fox Village, Alaska, in 1899

For transportation, Harriman had a steamship, the SS George W. Elder, retrofitted with luxurious staterooms for the members of the expedition, a stable for animals, studios for taxidermists and artists, and even a library of more than 500 books about Alaska. The expedition left Seattle to much fanfare on 31 May, heading north up the coast of British Columbia to Alaska.

Over the next two months they explored the coasts and islands of Alaska as far north as the Bering Strait where they landed on both the Alaskan and Russian shores. Sometimes expedition parties went ashore to camp, hunt, paint, take photographs and collect specimens. Most stops were short as Harriman relentlessly wanted to press on northward. In all, they travelled 14,500 km (9000 miles) by ship, returning to Seattle on 30 July.

Routes and stop in Alaska. Map by Fuertes

During the course of the expedition, thousands of specimens were collected, a new fjord discovered, 5000 photographs, taken and hundreds of paintings and illustrations produced. Though only 25 years old, Fuertes was already at the peak of his powers as a bird artist and illustrator:

A few of Fuertes’ illustrations from the expedition

On return Harriman funded the production of 14 volumes of scientific observations, well-illustrated with photographs, drawings and colour plates of paintings. The section on birds was written by Charles Keeler and is largely a narrative of what they saw at different stopping points. Though the expedition did not produce anything very interesting from an ornithological perspective, it was not bad, as Harriman himself said: “as a summer cruise for the pleasure and recreation of my family and a few friends”.

SOURCES

  • Burroughs J, Muir J, Grinnell GB (1901). Alaska; Narrative, glaciers, natives; Volume I. New York: Doubleday, Page & Company
  • Keeler C (1902) Days among Alaska birds,. pp 205 – 234 IN Dall W, Keeler C, Fernow BE, Gannett H, Brewer WH, Merriam CH, Grinnell GB, Washburn ML. History, geography, resources. Harriman Alaska Series, Volume 2. New York: Doubleday, Page & Company
  • various authors (1901- ) Harriman Alaska Series, Volumes 1-14. New York: Doubleday, Page & Company
  • websites: Biodiversity Heritage Library, Wikipedia, PBS, Cornell University, Smithsonian

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:

Grew2

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)

SOURCES

  • 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.

Footnotes

  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

Dirty Birds

BY: Bob Montgomerie, Queen’s University | 4 February 2019

As a teenager, in the 1960s, I spent much of my spare time during the school year hanging out at the Royal Ontario Museum (ROM), not far from my family’s home in Toronto.  The denizens of the bird/mammal prep room in the basement of the museum—Don Baldwin, Hisham Badran, Jim Borack and Rolph Davis—took me under their wing and taught me how to prepare bird skins and skeletons for the collections. That experience led directly to my Dream Jobs #3 and #4 in 1967/68 when I stopped going to the University of Toronto in search of a more satisfying (and financially rewarding) way to spend my days.

Dream Job #3 had me working as a preparator for the Mammalogy Department, mainly preparing bat skulls sent in by a collector from what we still called British Guiana as it had only been renamed Guyana a year before. My Dream Job #4 was a Park Naturalist position at Algonquin Park where my first duties were to put their museum’s specimen collection in order. The contrast between breathing formaldehyde all day in the museum lab and inhaling the intoxicating scent of piney woods reminded me that I was really a naturalist at heart.

The Ornithology Department at the ROM was thrown into a bit of a tizzy in 1965 when they hired Jon Barlow [1], with his newly-minted PhD, to be curator. Jon brought new ideas and new methodologies with him and those changes—not surprisingly—upset the old guard and intrigued the young. Jon introduced us all to quantification (statistics!), cladistics, and phenetics [2] for the purpose of classification; to the value of skeletal and tissue material for museum collections (and, of course, systematics); to the need for rigour in field notes and detailed specimen labels [3]; and to the notion that examining intraspecific variation was useful—essential even—for understanding evolutionary change and speciation.

For those of us in the prep room, the biggest changes were that we focussed more on skeletal and tissue collections, and that skins should be washed in detergent to remove blood and dirt, then dried before making them into a study specimen. Barlow also taught me the “John Williams method” [4] of skinning birds by making an incision on the side rather than the belly, stuffing the skin with a cone of cotton, and finishing up without stitching up that initial incision. I used that method for a while but never perfected it and to this day use the traditional method of preparing study skins as outlined in Rudolph Anderson’s classic book on preserving vertebrate animals.

bichicken
dirty Sacred Ibis in Australia

Barlow wanted us to wash bird skins not for aesthetic reasons, as you might expect, but because he was interested in measuring plumage colours accurately. Such colours would, of course, be obscured by dirt from dust-bathing or simply by the bird living in dirty environments, by foraging in evergreens where they would get resin on their feathers, or by blood and guts on their plumage while they were eating. He was interested in geographic variation in House Sparrows, to see how their colours may have varied (evolved) as they spread across the continent since their introduction in New York in 1851/52. He had studied with Richard Johnston at the University of Kansas and wanted to contribute to Johnston’s evolutionary analyses of that species. At first, we measured plumage colours by simple comparison to colour swatches in the Villalobos Atlas de los colores [5]. But Barlow was interested in exact quantification so we soon tried to modify a cumbersome desktop transmission spectrophotometer to measure plumage reflectance.

A recent paper by Shane DuBay and Carl Fuldner, at the University of Chicago, now suggests that by washing those bird skins we were flushing down the sink an important record of atmospheric pollution. On noticing that some bird specimens at the Field Museum of Natural History were quite dirty looking, they decided to investigate the source of that dirt. It turned out to be largely black carbon, which they verified by examining feather under a scanning electron microscope. They then developed a method using digital photographs of birds with white breasts to quantify the degree of light reflectance (less reflectance = more carbon deposited).

2ETOLIGY5A5YXESY2PALA36WVA
Horned Larks when atmospheric carbon was high (left) and today (right

To examine variation in these carbon deposits on birds in the industrial heartland [6] of America, the researchers photographed and analyzed the white breasts of 1097 specimens of five species [7] collected over the past 135 years. They used the excellent collections at the Field Museum, the Carnegie Museum of Natural History, and the Michigan Museum of Zoology. Because these species all moult in the fall, and every fall, they took specimens only from the winter months, and could thus date stamp the deposition of carbon on the plumage to within a few months. The remarkable results are shown on the graph below. The black line shows their estimate of average carbon accumulation on the specimens from 1888-2015; the orange line shows the quantity of coal burned in the USA during that period; and the purple line shows the results of a comprehensive model to predict the black carbon emissions in those states.

Using the birds as an index of black carbon in the atmosphere, it looks like the amount of carbon was highest from 1880 until the beginning of the Great Depression (red arrow), when it declined steeply as carbon emissions dropped. Emissions rose again as the depression ended and WWII began (blue arrow), then declined slowly to its present level after the Air Pollution Control Act (green arrow) and other regulations were imposed to reduce emissions even though consumption was on the rise (orange line).

CARBONgraph
graph modified from DuBay and Fuldner (2017)

Why are these results important? For one thing they suggest that the previous model of atmospheric carbon made an estimate that was too low from 1880-1910. This is an valuable bit of knowledge for climate change models. The results also match the estimate of peak atmospheric carbon in 1906-1910 as estimated from a Greenland Ice core. That core presumably measured what was in the atmosphere over Greenland, but this new study corroborates that finding. It would be interesting now to compare the sootiness of birds from this study to birds collected outside those industrial states, and even in other parts of the world.

Museums worldwide have suffered from shortages of funds and staff since at least the 1970s. Just like blue-skies research, the essential role of museums for science and society is hard to quantify. The historical value of museums is obvious and they are clearly invaluable for systematics research. But we can only guess what new discoveries will emerge from museum collections and they deserve our support. I would never have imagined that they might help us to understand climate change.

SOURCES

  • Anderson RM (1932) Methods of collecting and preserving vertebrate animals. Ottawa: King’s Printer.
  • DuBay SG, Fuldner CC (2017) Bird specimens track 135 years of atmospheric black carbon and environmental policy. Proceedings of the National Academy of Sciences (USA) 43: 11321-11328
  • Johnston RF (1973) Evolution in the House Sparrow, IV. Replicate studies in phenetic covariation. Systematic Zoology 22: 219-226.
  • Johnston RF,  Selander RK (1971). Evolution in the House Sparrow, II. Adaptive differentiation in North American populations. Evolution 25:1-28.
  • Ridgway RA (1912) Color standards and color nomenclature. Washington, DC.

  • Villalobos-Dominguez C, Villalobos J (1947) Atlas de los colores. Buenos Aires: Libreria El Ateneo Editorial.

Footnotes

  1. Jon Barlow: (1935-2009) was the first person with a PhD in ornithology to be Curator of Ornithology at the ROM. He was curator there for 35 years.
  2. (statistics!), cladistics, and phenetics: Barlow did his PhD at the University of Kansas where he learned about all of these topics from Robert R. Sokal who wrote classic books on both biostatistics and numerical taxonomy
  3. detailed specimen labels: Barlow wanted so much info on the labels (life size drawings of gonads and skull ossification, preparation methods, details of moult, tissue and skeleton specimens) that we often had to use 2-3 labels to contain it all.
  4. John Williams method: Williams (1913-1997) was well-known as an expert on African birds and was said to be able to prepare >30 high quality bird skins a day using his method. My record was more like 15 on my very best day.
  5. Villalobos Atlas de los colores: there were other colour atlases more widely used by ornithologists (see here), like Ridgway (1912) but Jim Baillie (see here), the acting curator before Barlow arrived, had noticed a stack of the Villalobos atlases remaindered in a local bookstore and snapped them up at $5 each. The Villalobos had lots of colour swatches finely grading into one another, and clever little holes in each swatch so you could more easily match them to the object of interest.
  6. industrial heartland: Pennsylvania, Ohio, Indiana, Michigan, Illinois, Wisconsin
  7. five species: Field and Grasshopper Sparrows, Eastern Towhee, Horned Lark, and Red-headed Woodpecker

Bird Paper One

BY: Bob Montgomerie, Queen’s University | 10 December 2018

When we were writing our Ten Thousand Birds book on the history of ornithology since Darwin, we thought it might be interesting to try to illustrate the growth of the field since the mid-1800s. To do that, we prepared a graph showing the number of articles and books published per year for every fifth year since 1865, using both Zoological Record and, for recent years, Google Scholar. The results were staggering [1], showing an explosive growth in publications on—and presumably knowledge about—birds since the second world war. Since the year 2000, there have been more articles and books published about birds than in the entire period from the beginning of scientific publishing in 1665 until 2000. We can estimate the number of publications before 1865 with some confidence as there were very few bird papers published before that date. The world’s major bird journals did not even start publishing until the mid-to-late 1800s [2].

Untitled

When we compiled that graph, we wondered when the first-ever scientific paper had been published on birds. It had to be after 1664, as the first ever scientific journals [3], Journal des sçavans and Philosophical Transactions, began publishing early in 1665.

To find that first bird paper, I scoured the early issues of both journals, looking at each issue as there was no Zoological Record or Google Scholar coverage that far back. The early issues of Journal des sçavans were devoted largely to obituaries, astronomy, and Cartesian philosophy, and Philosophical Transactions focused mainly on optics, astronomy, and other physical phenomena in its earliest years, though most issues had at least one paper on a biological/medical topic [4].

Bee_hummingbird_(Mellisuga_helenae)_adult_male_in_flight
Male Bee Hummingbird

Although birds were mentioned in several papers in the first few years of scientific publication, the first paper exclusively about birds did not appear until May 1693—in the 17th volume, and 200th issue, of Philosophical Transactions. That paper was attributed to the noted English botanist Nehemiah Grew [5] who published a letter (by a Mr Hamersly [6]) describing a hummingbird. He called the bird both ‘Hum Bird’ and ‘Tomineius’, the latter a Spanish word derived from ‘tomino‘ which was a measure of weight equal to 12 grains (0.78 g). In his Ornithology of Francis Willughby, published in 1678, John Ray suggests that the name ‘Tomineius‘ reflects the weight of the bird. But the smallest hummingbird—the Bee Hummingbird (Mellisuga helenae) of Cuba—weighs three times that. I suspect that the name ‘Tomineius’ was just meant to indicate that the bird was extremely small.

Antillean-Crested-Hummingbird-Orthorhyncus-cristatusWe don’t know for sure which species Hamersly was referring to as tomineius was probably a general term for all hummingbird species. In his 1590  book on the West Indies, for example, José de Acosta says that hummingbirds were called ‘Tomineios‘ in Peru.

A hand-written annotation  in some manuscript notes [7], presumably by Hamersly, found in a copy of Richard Ligon’s 1657 book on Barbados says that “I sent this description of this bird to doctor Grew one of the Royal Society & he caused it to be printed in their philosophical transactions. This suggests that Hamersly was describing one of the 3 species that are common on Barbados. By ‘this bird’, the annotator was referring to the picture below right, which, though crude, looks most like the Green-throated Carib (Eulampis holosericeus. By its size, however, I think Hammersly must have been referring to the Antillean Crested Hummingbird (Orthorhyncus cristatus) which weighs about 3 grams, the smallest hummingbird on the island. It’s too bad that we don’t really know who Hamersly was, nor which species he was describing.

GtCarib
Green-throated Carib (L), and hummingbird illustration (R) from manuscript notes [7]
 

Grew, or rather Hamersly, made a number of perceptive observations of the hummingbirds, though recent research has shown that he was not quite correct. Here are a few of the interesting things that Hamersly noted, with comments and what we now know in square brackets:

  • “He is of a most excellent shining green Color…resemble some of our English Drake-heads” [true, both are iridescent green]
  • whole weight was the tenth part of an ounce Avoirdupoise” [this would be about 2.8 g which is about right for the Antillean Crested Hummingbird]
  • They feed by thrusting their Bill and Tongue into the Blossoms of Trees, and so suck the sweet juice of Honey from them” [hummingbirds don’t suck [8]; they take up nectar into their grooved tongue and force it back into their throat by pressing their tongue with their bill as it retracts]
  • I did observe them several years but never heard them sing” [he claims they don’t sing, but they do, as do all of the 50 or species that I know reasonably well. He may have meant they don’t sing a song that sounds like most of the passerine bird songs and that is generally correct]
  • He is called the Hum-bird or Humming Bird because some say he makes a noise like a Spinning Wheel when he flies..I never heard any Noise; besides their Body and Wings are too small to strike the Air to make any Noise” [he is mistaken here, of course, but he later acknowledges that other people have heard them humming. He should have known that mosquitoes make noise so that his comment about size must be wrong.]

He did correctly note that they are very solitary, and suggested that with such a beautiful plumage they may not need to sing well: “so I think this Bird is so beautiful to the Eye, as not at all to please the Ear“. Indeed, recent studies have found such a tradeoff between selection for elaborate song or bright plumage in different groups of bird [9].

In Nehemiah Grew’s day, anyone interested in the sciences could read everything published in all (both) of the scientific journals. Even when I was a PhD student, in the 1970s, it was possible (and de rigueur) to read most of the papers in ecology and evolution published in the major journals, and to read all of the recent papers published on your study organism. Those days are over and few scientists can manage to even be aware of all of the research relevant to their own studies. No wonder many scientists get most of their information about recent studies in their field from Twitter.

Even if you wanted to keep up with research on birds since 2000, you would face a daunting task. The Web of Science [10] says that 127,000 papers have been published on birds from 2000 to 2018. The following graphic shows the distribution of 115,000 of those papers in the best-studied topics:

ornithology
Papers on birds published 2000-2018 on the 10 most common topics

Even focussing on hummingbirds, you would have to read 2383 papers to be fully informed about research published since 2000 (see below). Contrast this to the 36 papers on hummingbird ecology and evolution published during my PhD years, and the 48 papers published on those topics from 1900 until the year I graduated in 1979. We have come along way since Grew began the scientific publications about birds.

hbird papers
Papers on hummingbirds published from 2000-2018 on the 10 most common topics

SOURCES

  • Badyaev AV, Hill GE, Weckworth BV (2002) Species divergence in sexually selected traits: increase in song elaboration is related to decrease in plumage ornamentation in finches. Evolution 56: 412–419

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

  • Boyle (1865) A way of preserving birds taken out of the egge, and other small fætus’s. Philosophical Transactions 1: 199-201

  • de Acosta J (1590) Historia Natural y Moral de las Indias. Sevilla: Juan de Leon.
  • 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
  • Lefanu W (1971) The Versatile Nehemiah Grew. Proceedings of the American Philosophical Society 115: 502-506
  • Ligon R (1657) A true & exact history of the island of Barbados: Illustrated with a mapp of the island, as also the principall trees and plants there, set forth in their due proportions and shapes, drawne out by their severall and respective scales. Together with the ingenio that makes the sugar, with the plots of the severall houses, roomes, and other places, that are used in the whole processe of sugar-making; viz. the grinding-room, the boyling-room, the filling-room, the curing-house, still-house, and furnaces; all cut in copper. London: Printed for Humphrey Moseley, at the Prince’s Arms in St. Paul’s Church-Yard.

  • Price DJS (1963) Little science, big science. New York: Columbia University Press.

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

  • Rico-Guevara A, Fan T-H, Rubega MA (2015) Hummingbird tongues are elastic micropumps. Proceedings of the Royal Society B: Biological Sciences 282:20151014.

  • Shutler D, Weatherhead PJ (1990) Targets of sexual selection: song and plumage of wood warblers. Evolution 44:1967–1977.


Footnotes

  1. staggering results: while the numbers are high, the pattern is typical of all sciences, as described by Derek da Solla Price in his 1963 book
  2. first major ornithological journals: see previous posts, here, here, and here
  3. first ever scientific journals: Journal des sçavans began publishing on 5 Jan 1665, and Philosophical Transactions (later called Philosophical Transactions of the Royal Society) on 6 March of that same year.
  4. one paper on biological topics: Philosophical Transactions did have early papers on snake behaviour, breeding silkworms, and various medical anomalies, for example. There was even an 1865 paper on preserving bird embryos (Boyle 1865) but I don’t really count that as being about birds
  5. Nehemia Grew: was an early Fellow of the Royal Society, and was both editor of Philosophical Transactions (1678-79) and secretary of the RS. He is often called the Father of Plant Anatomy
  6. Mr Hamersly: although Grew published this letter under his own name, he revealed in Grew (1693b) that it was actually written by a Mr Hamersly of Coventry. See here for more information, suggesting it might have been John Hamersly, also referred to here as John Hammersley.
  7. manuscript notes: see here for details
  8. hummingbirds don’t suck: I discovered this tin the 1970s when doing my PhD on hummingbirds but I am sure that all of the other biologists I knew who were studying hummingbirds in those days—Bill Calder, Peter Feinsinger, Lee Gass, Larry Wolf, Reed Hainsworth and Frank Gill—knew this too; Rico-Guevera et al. (2015) recently described the details and physics of this process
  9. tradeoff between song and plumage: see for example, Shutler and Weatherhead (1990) and Badyaev et al. (2002)
  10. Web of Science: These numbers are smaller than what we show in the graph at the top of this essay, because Web of Science focuses only on publications in scientific journals, whereas we graphed all publications about birds. The data from Web of Science show the same patterns as in that graph but only about 7000 bird papers published in 2010, for example. To generate those data I entered bird* and ornithology as topics for the first graph and hummingbird* for the second and searched through the years in question

IMAGES: top graph from Birkhead et al. (2014), drawn by the author; Bee Hummingbird from Wikipedia; stamp from a stamp collection website; Green-thoated Carib (on Barbados) photo from Wikipedia, painting from manuscript notes [7]; bottom two graphs from Web of Science (accessed 9 Dec 2018)

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.

Linnaeus
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.

GrinnellFieldNotes
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.

SOURCES

  • 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.


Footnotes

  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].

HudsonMAP
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.

clet
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]

Tuck52Digges
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.

SOURCES

    • 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.

    Footnotes

    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.

Much Ado About a Cockatoo (reposted)

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

For the past week or so the internet has been abuzz about a cockatoo depicted 4 times in the margins of Frederick II’s De Arte Venandi cum Avibus written around 1245 CE [1]. The story is that this bird suggests a mediaeval trade route from Australia to Italy, overturning the Eurocentric notion that Australia was a dark continent until ‘discovery’ by Dutch sailors early in the 17th century. This story has already been retweeted about 2000 times on Twitter, and has appeared in the popular press worldwide, including CNN, Reuters, ABC (Australia), Japan Times, and The Guardian [2].

CockatoosFII
Marginal cockatoos (numbers indicate folio pages)

This story began with an original paper published last month  in the journal Parergon [3] by Heather Dalton, Jukka Salo, Pekka Niemelä and Simo Örmä. That paper is wonderfully detailed about the creation and provenance of De Arte, about the source of Frederick’s cockatoo, and the details of the four coloured drawings of the cockatoo in the surviving copy of the original manuscript housed in the Vatican Library in Rome [4]. The popular press has been remarkably accurate in reporting the details of that paper, avoiding the hyperbole and small misleading errors that too often characterize science journalism.

Here, in a nutshell, are 5 quotes that summarize what I think are the important features of this story:

  1. Frederick_II_and_eagle
    Frederick II

    “Frederick II of Sicily made contact with the Kurdish al-Malik Muhammad al-Kamil in 1217… The two rulers communicated regularly over the following twenty years, exchanging letters, books and rare and exotic animals….[like] the Sulphur-crested or Yellow-crested Cockatoo the sultan sent Frederick.”  [5]

  2. “De arte was written in Latin by Frederick or a scribe under his direction between 1241 and 1244…Amongst the nine hundred marginal illustrations of birds, animals, falconers, perches, and falconry equipment are four coloured drawings of the white cockatoo gifted to Frederick II.” [5]
  3. “Discovery of earliest European depiction of cockatoo in medieval book rewrites history of global trade” [6] see also quote 7
  4. “Because the four images in the Vatican manuscript have rarely been reproduced in print, few people are aware of their existence. This may be because many scholars have relied on Casey Albert Wood and Florence Marjorie Fyfe’s 1943 English translation of all six books of the De arte. 11 Although Wood and Fyfe included many illustrations from the Codex Ms. Pal. Lat. 1071, they did not include those of the cockatoo. [5]
  5. “Bearing in mind the shape of the crest, the blue/grey of the periophthalmic ring and the lack of a yellow tinged ear patch, Frederick’s cockatoo was in all likelihood a Triton Cockatoo…or one of the three subspecies of Yellow-crested Cockatoos that have a yellow crest.” [5]
  6. “The main significance about it is we tend to think of our region, not just Australia, but the islands around it, as the very last things to be discovered; the European view is it’s almost this dead continent and nothing was happening until Europeans discovered it.” [7]

While Dalton and colleagues have done a great job summarizing all of the details in their paper, I do have a few quibbles with the final four points listed above. I was, for example, a little surprised to hear of the ‘discovery’ of these cockatoo drawings because I certainly knew about them. As so often happens, I wondered if I had simply failed to realize their significance.

But as the paper so nicely summarizes, previous authors [8] had written about the cockatoos, so this new work might be better characterized as a re-discovery. In fairness, Dalton and colleagues never claimed this to be a new discovery but this is the way that most of the popular press has characterized their work.

WoodFyfeP38
Wood and Fyfe p 38: cockatoo is at top of right margin

They do claim (point 4), however, that modern scholars are generally unaware of these illustrations, largely because everyone reads Wood and Fyfe’s translation from 1943. But that’s how I knew about the cockatoo—one of the pages (folio 18v) with the cockatoo in the margin is reproduced, albeit in black and white, on page 38 in Wood and Fyfe’s book. In the caption they even say  “also containing the reference to the parrot (?) sent to Frederick by the Sultan of Babylon”, and in a footnote on page 59 they say it was likely a cockatoo from the Sunda Islands.

Dalton and colleagues do a really nice job of describing the four coloured marginal drawings and they use those details to try to identify the bird. They make the reasonable conclusion that it was likely a female Sulphur- or Yellow-crested Cockatoo. Others have suggested that it might be a White Cockatoo [8]. Nonetheless, I cannot agree with the authors’ assumption that those descriptions are very useful for identification.

cockatoosBoW

For example, in the De Arte illustrations, it looks like the birds have yellowish flanks and back, whereas none of the potential cockatoos have that colouring [9]. They also interpret the shape of the crest as ruling out the White Cockatoo, but the shape and size of the bill, feet, and wing feathers are so inaccurate that I would not be inclined to assume that the crest is correctly drawn. Moreover, none of the illustrations in De Arte show the characteristic yellow cheek patch of the Yellow- and Sulphur-crested.

peacockFiiThe marginal drawings are, after all, crude by modern standards, as you can see from the reproductions above. The manuscript has at least 7 marginal drawings of peacocks [10], for example, that show that, although the artist was remarkably good for his day, he was no Lars Jonsson. Thus it would be entirely reasonable for Frederick’s bird to be a White or Yellow-crested Cockatoo, from Sulawesi (one of the Sunda Islands) or the Moluccas [11]. If that is correct then I am not so sure that this cockatoo really tells us anything about mediaeval trade routes.

It has long been known that there was extensive trade between southeast Asia and the Middle East along the Silk Road, beginning centuries before Frederick’s day. That ‘road’ includes several marine routes extending as far east as Sulawesi and the Sunda Islands (see map below). While there may have been some trade between New Guinea and northern Australia in the 13th century, there is really no evidence for this, as far as I know. Thus, Australia does seem to have been a relatively ‘dark continent’ until the 17th century, especially to Europeans, and Frederick’s cockatoo does not really shed any light on that narrative.

I feel that I should emphasize how much I enjoyed the original article by Dalton and colleagues, despite my reservations above. In my view, science progresses when there is some healthy skepticism, and that’s what I have tried to present here.

Silk_route
Land and marine trade routes in mediaeval times.

NOTE: I accidentally posted an incomplete version of this essay a couple of days ago. My apologies for that. I blame the heat.

SOURCES

  • Dalton H, Salo J, Niemelä P, Örmä S (2018) Frederick II of Hohenstaufen’s Australasian Cockatoo: Symbol of Detente between East and West and Evidence of the Ayyubids’ Global Reach. Parergon 35: 35-60.

  • Frederick II (~1245) De arte venandi cum avibus. Vatican City, Biblioteca Apostolica Vaticana, codex Ms. Pal. Lat. 1071.
  • Kinzelbach R (2008) Modi auium – Die Vogelarten im Falkenbuch des Kaisers Friedrich II’. pp 62–135 in Vol 2 of Kaisers Friedrich II 1194–1250: Welt und Kultur des Mittelmeerraums (ed. Ermete K, Mamoun Fansa M, and Carsten Ritzau C). Mainz: Philipp von Zabern.
  • Rowley, I. (2018). Cockatoos (Cacatuidae). In: del Hoyo, J., Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (eds.). Handbook of the Birds of the World Alive. Lynx Edicions, Barcelona. (retrieved from https://www.hbw.com/node/52255 on 2 July 2018).
  • Stresemann E (1975) Ornithology from Aristotle to the present. Cambridge, MA: Harvard University Press.

  • Willemsen CA (1980) Das Falkenbuch Kaiser Friedrichs II. Nach der Prachthandschrift in der Vatikanischen Bibliothek. Dortmund: Harenberg.

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

  • Yapp WB (1983) The illustrations of birds in the Vatican manuscript of De arte venandi cum avibus of Frederick II. Annals of Science 40: 597–634

Footnotes

  1. Frederick II’s manuscript: see my earlier post on this ornithologically important masterpiece here
  2. articles in popular press: but curiously not (yet) The New York Times or any of the other leading American and Canadian news media outlets.
  3. Parergon: is the journal of the Australian and New Zealand Association of Medieval and Early Modern Studies (Inc.), and has been publishing refereed articles since 1983.
  4. Frederick’s manuscript in the Vatican Library: is also available for study online in two spectacularly reproduced digital copies here and here.
  5. Quotations 1, 2, 4 & 5: from Dalton et al. 2018
  6. Quotation 3: from the headline in The Telegraph (UK)
  7. Quotation 6: Helen Dalton quoted in The Guardian (UK)
  8. previous authors on this cockatoo: see, for example, Wood and Fyfe (1943), Stresemann (1975), Willemsen (1980), Yapp (1983), Kinzelbach (2008)
  9. yellowish back and flanks: I would be inclined to interpret this colour as shading rather than as the colour of the feathers.
  10. peacocks: would probably have been a very familiar bird in the courtyards of Italy in the 13th century, having been traded along the Silk Road for centuries before. Like the cockatoo, those drawings would have been based on live speciemens
  11. likely species: Dalton et al. (2018) do seem to favour the Yellow-crested Cockatoo in their analysis, so it is curious to me that they make such a fuss about the trade routes, since the range of the Yellow-crested on one of those mediaeval routes

IMAGES: Cockatoos and peacocks from De Arte copied from the online versions; Frederick II and Silk Road map from Wikipedia; Cockatoos and range map from Handbook of Birds of the World (Rowley 2018); page from Wood and Fyfe scanned from the author’s copy.

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.

Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997
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]

GouldGSWoopecker
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.

SOURCES

  • 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.


Footnotes

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

Audubon’s Legendary Experiments

BY: Bob Montgomerie, Queen’s University | 11 June 2018

In his Ornithological Biography [1], John James Audubon describes two experiments that have become legends in the annals of ornithology. Both were ingenious ideas, but Audubon’s conclusions were misleading and remained uncorrected for more than a century. According to  a new paper by Matthew Halley in Archives of Natural History, one of those experiments might never have been conducted at all.

mill grove
Audubon mansion at Mill Grove

In the spring of 1804, just before his 19th birthday [2], Audubon found an Eastern Phoebe nest in a cave near where he was living, at the plantation his father had purchased at Mill Grove, Pennsylvania [3]. Audubon had arrived from France a few months earlier and immediately began exploring the countryside where, he said “Hunting, fishing, drawing, and music occupied my every moment; cares I knew not, and cared naught about them. I purchased excellent and beautiful horses, visited all such neighbors as I found congenial spirits, and was as happy as happy could be.” [4]

Audubon wondered whether his phoebes might exhibit what we would now call ‘natal philopatry’: “I had at that period an idea that the whole of these birds were descended from the same stock.” [5]. To test that idea, he put some ‘light threads’ on the legs of the 5 nestlings. The birds, or their parents, repeatedly removed those threads but finally gave up [6]. Once the birds had habituated to the threads, Audubon says that “when they were about to leave the nest, I fixed a light silver thread to the leg of each, loose enough not to hurt the part, but so fastened that no exertions of theirs could remove it.” [7]. Others have, I think, misinterpreted Audubon’s ‘silver thread’ as being silver-coloured yarn but the term is (and was) also used for wire made of silver. Audubon later refers to these as ‘rings’ (see below) suggesting that they were solid and unlikely to be removable by the birds.

1863_17_120_EasternPhoebe_OE

The next year, Audubon claims, he found two of those marked phoebes—which he called pewees—nesting near where they were born:

…I had ample proof afterwards that the brood of young Pewees, raised in the cave, returned the following spring, and established themselves farther up on the creek…At the season when the Pewee returns to Pennsylvania, I had the satisfaction to observe those of the cave in and about it. There again, in the very same nest, two broods were raised. I found several Pewees nests at some distance up the creek, particularly under a bridge, and several others in the adjoining meadows, attached to the inner part of sheds erected for the protection of hay and grain. Having caught several of these birds on the nest, I had the pleasure of finding that two of them had the little ring on the leg.” [8]

Now Halley—correctly I think—calls into question the veracity of these observations of natal philopatry, First, he points out, the 40% return rate (2 of 5 banded) of Audubon’s nestling phoebes far exceeds the rate of natal philopatry for this species—and indeed all passerine birds [9]—revealed in subsequent research with large sample sizes. Work done from 1988-90 at my own institution’s field station by Kelvin Conrad, for example, showed a 1.3% return rate of 217 banded nestlings [10]. A much larger study in Indiana [11] found that only 218 of 11,847 (1.8%) phoebe nestlings returned to anywhere within a 250-kmstudy area. Based on those numbers, the chance that 2 of Audubon’s 5 nestlings actually returned is almost zero.

Second, Halley checked the dates of Audubon’s return to France (12 March 1805) and subsequent arrival back in Mill Grove (4 June 1806). Based on the normal first egg dates from nearby sites, Halley argues that Audubon could not have been at Mill Grove during the 1805 breeding season and could therefore not have observed the returns he claimed. While I think Halley is correct, it is always possible that Audubon was mistaken about the years when he conducted this experiment.

My guess, though, is that Audubon did actually make up this story, not about banding the birds [12] but about them returning to their natal site. Audubon was well known to be desperate to establish his reputation as an ornithologist, in part, at least, to enhance the sales of his fabulous collection of etchings based on his paintings of North American birds. Audubon was often at odds with Alexander Wilson and George Ord, sometimes plagiarizing Wilson and fabricating evidence so that he would be seen as the first to record a species of bird or an interesting observation [13]. Elliott Coues, for example, said that Audubon: “..loved warmth, color, action; he liked to exaggerate and ’embroider,’ and make his page glow like a hummingbird’s throat, or like on of his many marvellous pictures; he had no genius for accuracy, no taste for dull, dry detail, no care for a specimen after he had drawn it.” [14]

Audubon’s other misleading experiment was designed to assess the ability of Black Vultures to detect odours, but that is a story for another day. Whether or not Audubon was always accurate in his descriptions of bird behaviour or his timelines of events, his experiments were certainly ingenious. He probably does deserve credit as the first person to band birds in North America, and he really could paint a marvellous picture:

black-vulture-or-carrion-crow-john-james-audubon
Audubon’s Black Vultures

SOURCES

  • Audubon JJ (1831-1839) Ornithological biography : or an account of the habits of the birds of the United States of America; accompanied by descriptions of the objects represented in the work entitled The Birds of America, and interspersed with delineations of American scenery and manners.  5 volumes. Edinburgh: Adam & Charles Black

  • Audubon JJ (1827-1838) The Birds of America. Edinburgh & London: J. J. Audubon.

  • Audubon MR (1897) Audubon and His Journals, with Zoological and Other Notes by Elliott Coues, vol. 1. New York: Charles Scribner’s Sons.

  • Burns FL (1908) Alexander Wilson: II. The Mystery of the Small-Headed Flycatcher. The Wilson Bulletin 20:63–79.

  • Halley MR (2018) Audubon’s famous banding experiment: fact or fiction. Archives of natural history 45:118–121.

  • Weatherhead PJ, Forbes MRL (1994) Natal philopatry and the cost of dispersal in Passerine birds. Behavioural Ecology 5:426–433.

  • Weeks Jr HP (2011) Eastern Phoebe (Sayornis phoebe), version 2.0. In The Birds of North America (A. F. Poole, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA.

Footnotes

  1. Ornithological Biography: designed by Audubon, with the help of William MacGillivray, to accompany his paintings in Birds of North America. This 5-volume treatise (available here) contains many interesting details and insights about birds but is not an easy read.
  2. 19th birthday: Audubon was born on 26 April 1785 in the French colony that is now Haiti
  3. Audubon’s home: Audubon travelled from France to Mill Grove by himself, on the orders of his father to manage the plantation but was more interested in natural history and he eventually drove the plantation into bankruptcy
  4. quotation about hunting etc: from Audubon 1897, page ; one of the neighbours he talks about was William Blakewell, father of the woman he married a couple of years later
  5. quotation about phoebe philopatry: from Audubon (1831-39) vol 2, 1834, page 125
  6. phoebes removing threads: this thread removal and habituation must have occurred over less than two weeks as phoebes leave the nest after 16-18 days
  7. quotation about silver threads: from Audubon (1831-39) vol 2, 1834, page 125
  8. quotation about returning phoebes: from Audubon (1831-39) vol 2, 1834, page 125-127
  9. natal philopatry of passerines: Weatherhead and Forbes (1994) summarized the rates of natal philopatry for 32 migratory passerines from various published and unpublished studies. The highest recorded rate of natal philopatry was 13.5%, and the majority of species had rates <5%, much lower than Audubon recorded for his phoebes
  10. natal philopatry in Ontario: unpublished data summarized in Weatherhead and Forbes (1994)
  11. natal philopatry in Indiana: unpublished data summarized in Weeks (2011)
  12. Audubon banding birds: I am inclined to believe that Audubon did band those birds as the details ring true. He thus deserves his reputation as America’s first bander, preceding by almost a century the next attempt to band birds in the America’s
  13. Audubon’s conflicts and plagiarism: see Halley (2018) for a brief summary
  14. Elliott Coues quotation: as quoted by Burns 1908 page 68

IMAGES: Audubon mansion from Audubon (1897);  Audubon’s paintings from his Birds of America (1827-38)

George Ord’s warbler

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

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

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

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

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

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

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

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

Gmelin1789
top of page 985 in Gmelin (1789) describing Motacilla tigrina

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

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

Edwards1758
Edward’s (1758) Spotted Yellow Fly-catcher (L) and Yellow-tailed Flycatcher (R)

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

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

SOURCES

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

Footnotes

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

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