Ladies, Parakeets, and the Biogeography of an Extinct Bird

BY: Bob Montgomerie, Queen’s University | 25 June 2018

In 1850, an anonymous author published a superb diary of natural history observations called ‘Rural Hours by a Lady’ based on two years of exploring the woods and fields near Cooperstown, New York. The book was wildly popular, and it was not long before the author was revealed to be Susan Fenimore Cooper [1]. On page 146 she says:

Parakeet2It is well known that we have in the southern parts of the country a member of the Parrot tribe, the Carolina Parakeet. It is a handsome bird, and interesting from being the only one of its family met with in a temperate climate of the Northern Hemisphere. They are found in great numbers as far north as Virginia, on the Atlantic coast; beyond the Alleghanies, they spread themselves much farther to the northward, being frequent on the banks of the Ohio, and in the neighborhood of St. Louis. They are even found along the Illinois, nearly as far north as the shores of Lake Michigan. They fly in flocks, noisy and restless, like all their brethren…In the Southern States their flesh is eaten…Birds are frequently carried about against their will by gales of wind; the Stormy Petrels, for instance, thoroughly aquatic as they are, have been found, occasionally, far inland. And in the same way we must account for the visit of the Parakeets to the worthy Knickerbockers about Albany.  [2]

Here, she correctly describes the bird as being most common in the southeastern states, though seen regularly as far north as the Great Lakes west of the Allegheny Mountains. What she did not know, of course, was that these were two subspecies, with different morphologies, ecologies and migratory strategies, as described below.

Live_captive_Carolina_parakeet
Shufeldt’s photo

The Carolina Parakeet was still abundant throughout its range in 1850 but, like the Passenger Pigeon, was soon to be extirpated. The second-last individual was a female called ‘Lady Jane’ who died at the Cincinnati Zoo in 1917; the last of its kind being Lady Jane’s mate, a male named ‘Incas’ who died there in 1918, one hundred years ago. Coincidentally, Incas died in the same cage where Martha, the last passenger pigeon, had died in 1914 [3]. There were reports of sightings in the wild for another 40 years or so, in Florida and Georgia, but none of those records were authenticated. Among the North American birds that have become extinct since the arrival of Europeans, the biology of the breeding biology Carolina Parakeet may be the poorest known [4]. And there is, surprisingly, only one photo of the bird in a natural-looking setting [5], taken by the irrepressible Robert Shufeldt in about 1900.

 

A recent pair of papers by Kevin Burgio and colleagues uses all of the known specimens and sightings of this bird to reveal some interesting insights into its distribution, ecology, and taxonomy. There were 401 of those sightings recorded between 1564 and 1944, and nearly 800 specimens in museums and private collections worldwide [6], almost all collected in the 1800s. As shown on the graphs below, the number of records climbed exponentially from 1500 to 1900, reflecting the increases in exploring the new continent, in writing about natural history, and in preserving ornithological data and specimens. There was an uptick in collecting, or at least preserving specimens, from 1870-1900 when it became clear that the bird was disappearing [6].

graphs
Records and specimens with known dates—note the log scale on upper two graphs.

Analyzing records only from states where the parakeet was known to breed, Burgio and colleagues, georeferenced all the data and used 147 unique localities to create the species breeding distribution models shown on the map to the right below. The map on the left was produced in 1889 by Edwin Hasbrouck with the known range in his day (black shading) nicely matching the newly reconstructed ranges of the two subspecies.

Burgio and colleagues’ research also suggested (i) that the breeding range of this species was much smaller than previously thought, (ii) that the two subspecies, previously only vaguely defined by size and colour, actually had disjunct ranges and occupied somewhat different climatic niches, and (iii) that the western subspecies was almost certainly migratory where the eastern one was not.

The authors also hoped their analysis would help to inform current conservation practices in an effort to save the 8% of bird species currently threatened to disappear as a result of climate change. Parrots, in particular, are in bad shape, with 42% of species listed as threatened or endangered.

MAPS
LEFT from Hasbrouck (1889) estimating the limits of the parakeet’s historical range (black line) with shading showing the range in the 1880s. RIGHT from Burgio et al. (2017) estimating the ancestral breeding ranges of the two subspecies (Hasbrouck’s range limit shown as a red line)

 

pipe
Clay pipe ca. 1650

As much as I like those recent papers, I think it’s unfortunate that many biogeographers draw their maps as if animals obeyed political boundaries, as on the right-hand but not the left-hand maps above. The right-hand graph implies, for example, that the bird could never have crossed the US-Canada border as there was nowhere to go. Despite that, there is some evidence that it did occasionally occur in southwestern Ontario, possibly blown off course as Susan Cooper suggested above. At an archaeological dig at Grimsby, Ontario, for example, Walter Kenyon found a clay pipe that looks distinctly like a parrot, made by native peoples in the mid-1600s. And Rosemary Prevec found 3 Carolina Parakeet bones at a native site near London, Ontario, dated at around 1100 CE. Both of these findings are no more than suggestive and could have been obtained in trade with natives living further south.

Possibly more convincing are some observations that Samuel de Champlain recorded in his notes in 1615, in the woods near where I live in Kingston. He says that he  “…penetrated so far into the woods in pursuit of a certain bird which seemed to be peculiar, with a beak almost like that of a parrot, as big as a hen, yellow all over, except for its red head and blue wings, which made successive flights like a partridge.” [8] There are definitely no other birds even remotely resembling that description in eastern Ontario today.

fancyfeathers
Catalog page, New York Millinery and Supply Company, Inc., New York

None of this nationalism is really important to our understanding of the bird’s ecology and demise, except to note that at one time the species was clearly widespread and mobile. What is important is an attempt to understand why they went extinct, as even by the middle of the 1800s it appeared to be declining in numbers [6].

Burgio and colleagues point to habitat destruction and hunting as the likely causes. Not surprisingly, the parakeet’s feathers were prized for the millinery trade, with some reports suggesting that ladies hats were sometimes adorned with skins of the whole bird. The 1901 ad to the right, for example, shows a whole parrot (skin) in the lower right corner, for the bargain price of 25¢ a bird or $2.95 a dozen (about $7.50 and $88 in today’s currency). While the documentation is sketchy, it is also likely that this species was a popular cage bird in Germany as well as in North America. The only other known photo, besides Shufeltdt’s, is also one of a pet called  ‘Doodles‘, kept by Smithsonian malacologist Paul Bartsch. In 1900, ‘doodle‘ meant ‘fool‘ and not the ‘absentminded scribble‘, Google commemorative, or online scheduler that it is today. And I wonder if Bratsch gave it that name to reminder him what fools we are when let any species go extinct.

SOURCES

  • Anonymous [Cooper, SF] (1850) Rural Hours by a Lady. New York: G. Putnam.
  • Burgio KR, Carlson CJ, Tingley MW (2017) Lazarus ecology: Recovering the distribution and migratory patterns of the extinct Carolina parakeet. Ecology and Evolution 7:5467–5475.
  • Burgio K, Carlson C, Bond A (2018) Georeferenced sighting and specimen occurrence data of the extinct Carolina Parakeet (Conuropsis carolinensis) from 1564-1944. Biodiversity Data Journal 6:e25280.
  • Cokinos C (2000) Hope Is the Thing With Feathers: A Personal Chronicle of Vanished Birds. New York: Penguin.
  • Fuller E (2013) Lost Animals: Extinction and the Photographic Record. London: Bloomsbury.

  • Greene WT, Dutton FGFG, Fawcett B, Lydon AF (1883) Parrots in Captivity, v. 2. London: George Bell and Sons.

  • Hahn P (1963) Where is that Vanished Bird? Toronto: University of Toronto Press. [see this previous post for more on this book[
  • Kennedy CC (1984) Did Champlain stalk a Carolina Parakeet in southern Ontario in 1615? Arch Notes 84:55–62.

  • McKinley, D. (1960) The Carolina parakeet in pioneer Missouri. The Wilson Bulletin 72:274–287.
  • McKinley D (1977) Climatic relations, seasonal mobility, and hibernation in the Carolina Parakeet. Jack-Pine Warbler 55:107–124.
  • Prevec R (1984) The Carolina Parakeet—its first appearance in southern Ontario. Arch Notes 84:51-54.
  • Snyder NFR (2004) The Carolina Parakeet: Glimpses of a Vanished Bird. Princeton, NJ: Princeton University Press.

Footnotes

  1. Susan Fenimore Cooper: was a superb naturalist, author, and artist whose work was overshadowed in more ways than one by that of her father, James. She deserves recognition and a separate essay on her own work, Stay tuned.
  2. Cooper quotation: from Anonymous 1850 page 146
  3. ‘Incas’ the parakeet: Like Martha, the last Passenger Pigeon, Incas was frozen and sent to the Smithsonian, but he was lost in transit (Fuller 2013)
  4. breeding biology poorly known: see Snyder (2004)
  5. Shufeldt’s photo: is one of a pair of pet birds that Shufeldt borrowed from his friend Edward Schmidt, and it took him hours to get it to sit still enough on a cocklebur to make a decent photo (Cokinos 2001). Both of Schmidt’s birds later died from chewing on the bars of their cage, possibly from lead paint poisoning (Fuller 2013)
  6. declining numbers by mid 1800s: see Hasbrouck (1889)
  7. records and specimens: see Hahn (1963), McKinley (1960, 1977) and Snyder (2004) for background
  8. Champlain quotation: from Kennedy 1984 page 55

IMAGES:  first parakeet is by Robert Ridgway from Baird et al. (1905); graphs by the author based on data in Burgio et al. (2017, supplement)—parakeet is an engraving by Benjamin Fawcett in Greene et al. (1883); maps from the original papers; clay pipe from Kennedy (1984); millinery ad from the Smithsonian collection

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.

San_francisco_fire_1906
The fires of April 2006, San Francisco
Oceanodroma.macrodactyla
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

Crew-of-the-Academy-1905-expedition-sonoma.edu_
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].

CalifAcadamyOfSciAug28-2008img0640
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

MayrNG
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.

SOURCES

  • 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.

Footnotes

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

Galápagos sojourn

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

26 February 2018

Mr Charles Darwin
Westminster Abbey
UK

My Dear Charles

Post_office-Floreana
Post Office Bay, Floreana

My apologies for not writing last Monday as I had suggested I might when I wrote to you on your birthday. We were still on the Santa Cruz II ‘steaming’ from Floreana to Baltra on Monday morning and there was no way yo get a message out. I thought of leaving a postcard for you in the barrel at Post Office Bay on Floreana but that might take months to get to you, or be stolen by a tourist.

We had a great visit to the Galápagos Islands, stopping on Baltra, Santa Cruz (Cerro Dragon and Puerto Ayora), Isabela (Punta Vincente Roca), Fernandina (Punta Espinoza), and Floreana (Punta Cormoran and Post Office Bay) to hike, snorkel and/or simply watch and photograph wildlife. I see from your Voyage of The Beagle that you, too, stopped on Charles Island (now called Floreana) and Albermarle (now Isabela), but you also went to Chatham Island (now San Cristobal) and James Island (now Santiago). Certainly, things have changed since you were in the islands in Sept-Oct 1835.

200902-charles-darwin-beagle-map

First, and maybe most strikingly, there are now a lot of people on the Galápagos Islands. There are now settlements on Santa Cruz, Baltra, San Cristobal, Floreana and Isabela, by far the largest being Puerto Ayora on Santa Cruz with maybe 25,000 inhabitants (though ‘officially’ 15,000). On top of that a staggering 225,000 people visited the islands in 2015, by boat or plane. The islands are now almost entirely a national park, so travel is restricted to 54 sites on land and 62 for diving in the surrounding ocean. Visitors are limited to about 4 hours per site and must be accompanied by a trained guide.

Fortunately, the places we visited (except Puerto Ayora and vicinity) seemed to be in a relatively pristine state with well-marked trails, no trash, and abundant wildlife close at hand. The birds and reptiles are still exceptionally tame and the waters clear and teeming with life.

DSCF4538
G. fortis with foot pox, on Baltra

The (your) finches were also common to abundant just about everywhere we went. They certainly have not been scared off by human developments as we saw them even inside the airport buildings on Baltra and throughout the town of Puerto Ayora. Even we seasoned ornithologists and birders found the species hard to distinguish on any given island so you are to be forgiven for not initially noticing the proliferation of finch species there. The downside of increased human traffic to the islands is that we saw a high incidence of foot pox in the finches on Baltra, and a parasitic nest fly (Philornis downsi) is now posing a serious threat to some finch populations [1]. The finches are so closely associated with humans in some places that there now signs posted to tell people not to feed the birds.

You will recall that John Gould identified 12 species of ‘Galápagos’ finches from your collections. There continues to be debate about how many finch species are actually on the islands, especially as we are now using new molecular tools to help distinguish evolutionarily stable populations that might be worth designating as distinct species. During the 20th century biologists often defined species as reproductively isolated populations (the ‘Biological Species Concept’) but that has proven to be difficult to test empirically and not always useful, in my opinion. At my count there are now at least a half dozen ways to define species and the debate continues in a lively (and I think very productive) fashion.

The Handbook of Birds of the World Online now lists 14 species of Geospiza, plus the Vegetarian Finch (Platyspiza crassirostris), the Grey Warbler-finch (Certhidea fusca), and the Green Warbler-finch (Certhidea olivacea) for a total of 17 species of Darwin’s Finches. I expect that DNA analysis will add to this total in the coming years.

Peter and Rosemary Grant also discovered an instance of speciation through hybridization of an immigrant male Geospiza conirostris from Española Island with a female resident Geospiza fortis on Daphne Major in 1981 [2]. The descendants of this pairing (the Big Bird Lineage, see below) have only mated with each other over the last 37 years. These birds are reproductively isolated from the resident population of G. fortis by their distinctive song. Odds are that this tiny population of the hybrid species will go extinct, but the documentation of this event has given us an insight into a form of speciation that you may not have anticipated, though it is likely to be quite rare.

2017_Princeton_Grants_Finch_BigBird_Familytree

Sadly, some of the tortoises that you recognized as being distinct species are now extinct due to hunting by sailors, collecting by museums, predation by introduced rats and cats, and habitat destruction by goats. It is estimated, for example, that 200,000 tortoises were taken from the islands before 1900. The tortoise (Chelonoidis abingdoni) from Abingdon Island (now Pinta) went extinct only 6 years ago when the last male (“Lonesome George”) died in captivity at the (relatively young) age of just over 100 years. George was preserved as a taxidermic mount and is now on display at the Charles Darwin Research Station in Puerto Ayora. Also now extinct is C. nigra from Floreana, which you saw and collected, was extinct by 1846 having been hunted mercilessly by sailors and the penal colony on that island; and C. phantastica from Narborough Island (now Fernandina) which is known only from a single specimen collected by Rollo Beck for the California Academy of Sciences in 1906. All of the other tortoise species are considered to be endangered or at least vulnerable with populations <10,000 each and some only in the 100s. There is hope, however, in restoring some of them by captive breeding and the eradication of predators.

DSCF4232
Chelonoidis porteri in the ‘wild’ on Santa Cruz Island

As you might expect, your name is intimately associated with the Galápagos with an island (formerly Culpepper Island) now named after you, as well as a research station and a hostel in Puerto Ayora, a tortoise (C. darwini), and, of course, those finches.

Yr obd srvt

Bob

SOURCES

  • Darwin C (1840) The zoology of the voyage of H.M.S. Beagle, under the command of Captain Fitzroy, R.N., during the years 1832 to 1836. London: Smith, Elder & Co.
  • Grant BR, Grant PR (2008) Fission and fusion of Darwin’s finches populations. Philosophical Transactions of the Royal Society B: Biological Sciences 363:2821–2829.
  • Kleindorfer S, Dudaniec RY (2006) Increasing prevalence of avian poxvirus in Darwin’s finches and its effect on male pairing success. Journal of Avian Biology 37:69–76.
  • Koop JAH, Kim PS, Knutie SA, Adler F, Clayton DH (2016) An introduced parasitic fly may lead to local extinction of Darwin’s finch populations. Journal of Applied Ecology 53: 511–518.

Footnotes

1. parasitic nest fly and foot pox: see Koop et al. 2016 on the fly and Kleindorfer and Dudaniek on the pox

2. speciation through hybridization: see Grant and Grant (2008)

IMAGES: Big Bird lineage from https://www.princeton.edu/news/2017/11/27/study-darwins-finches-reveals-new-species-can-develop-little-two-generations; Galapagos map from http://www.prairiefirenewspaper.com/2009/02/reflections-on-charles-darwin; Post Office Bay photo from Wikimedia Commons; all other photos by the author

Pigeon Coup

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

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

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

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

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

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

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

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

thumbs_our-team_02
Paul Hahn

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

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

ROM2013_13563_1
One of the Passenger Pigeon mounts at the Royal Ontario Museum

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

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

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

SOURCES

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

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

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


FOOTNOTES

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

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

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

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

5. Quotations from Hahn 1963:1.

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

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

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

IMAGES: ROM photos by Brian Boyle, courtesy of Mark Peck (both at the ROM); Paul Hahn from the Paul Hahn & Co. website at https://paulhahn.com/about/who-we-are/

The First Penguins

BY: Bob Montgomerie, Queen’s University | 23 Oct 2017

While preparing a talk [1] last week about the early history of ornithology in North America, I wondered who might have been the first to describe and identify a bird on this continent. As far as I can tell, that was Jacques Cartier when he wrote, in 1534, about the ‘Apponat‘ (originally in French but here in English translation [2]):

whose numbers are so great as to be incredible, unless one has seen them; for although the island is about a league in circumference, it is so exceedingly full of birds that one would think they had been stowed there Some of these birds are as large as geese, being black and white with beak like a crows. They are always in the water, not being able to fly in the air, inasmuch as they have only small wings about the size of half one‘s hand, with which however they move as quickly along the water as the other birds fly through the air. And these birds are so fat that it is marvellous. We call them apponatsand our two longboats were laden with them as with stones in less than half an hour. Of these, each of our ships salted four or five casks, not counting those we were able to eat fresh

Cartier also recorded seeing Margaulx (Gannets) and Godertz (probably Common Murres) but the apponat referred to here is the Great Auk (Puinguinis impennis). “Apponath” was what the local Newfoundland natives (i.e. Beothuks) called this bird. On his second trip to Isle des Oyseaux, a year later, Cartier wrote “The island is so exceedingly full of birds that all the ships of France might load a cargo of them without perceiving that any of them had been removed” [2].

The “isle of birds” (Isle des Oyseaux) referred to in that passage above, was called ‘Penguin Island’  in the 1600s and  was ‘officially’ called ‘Funk Island’ by the late 1700s. Cartier was not the first to visit this island, as Gaspar Corte-Real stopped there in 1501, and it is shown on two maps by Pedro Reinel—one in 1504 where he calls it ‘Y Dos Saues’ and the other in 1520 where it is labelled ‘Yihas das Aves’. The map below by John Mason was made around 1617 and clearly shows ‘Penguin Island’ off the northwest coast on Newfoundland. This map was drawn upside down (for some unknown reason), so Penguin Island is on the lower left margin.

mason-map-newfoundland-1617

There is some debate about where the word ‘penguin’ came from, though we can be certain that it was what Europeans called the Great Auk, centuries before any of the birds that we now call penguins had been ‘discovered’. The three most commonly suggested—but very different—origins for the word ‘penguin’, as applied to the Great Auk, are:

  1. Great_Auk_Thomas_Bewick_1804derived from the Welsh ‘pen gwyn‘, where ‘pen’ is their word for head (or headland) and ‘gwyn’ means white, referring either to the white patch on the bird’s head, or the fact that a headland full of Great Auks looks white. The Welsh (and other Europeans) would have known this bird long before they found it in North America, as it bred (and was slaughtered) across the eastern North Atlantic from Iceland thorough Great Britain and Norway to as far south as Spain. In 1577, Francis Fletcher, a clergyman who travelled with Sir Francis Drake, wrote in his log about the southern hemisphere penguins [3]: “Infinite were the Numbers of the foule, wch the Welsh men name Pengwin & Maglanus tearmed them Geese.” This seems to me fairly convincing evidence for the word’s origin.
  2. derived from the name ‘pin-winged’, referring to the lack of real wing feathers on this flightless bird. I like this explanation though the consensus seems to be that it is incorrect.
  3. derived from the Latin pinguis, meaning ‘fat’ or ‘plump’

There is quite a good online debate about these origins here, if you are interested, and the several books now published on the Great Auk variously mention and discuss where the word ‘penguin‘ came from.

Probably the first European to see what we now call penguins was the explorer Bartholomeu Diaz, from Portugal, who reached the Cape of Good Hope (Africa) in 1488, but he never mentioned these birds in his notes. The first mention of southern hemisphere penguins is probably in the travel book of Álvaro Velho who rounded the Cape of Good Hope with Vasco da Gama in 1497. He called them ‘otilicarios’ [opticians?] and said that (again in rough translation): “They are as big as ducks, but can’t fly because they have no feathers on their wings. These birds, of which we slaughtered as many as we could, cried like jackass.”

So the Great Auk was the first ‘penguin’. Presumably it seemed logical at the time to call the southern hemisphere flightless oceanic birds ‘penguins‘, as well, because they looked and behaved so much like the Great Auk. Hunted relentlessly, the Great Auk had disappeared from Funk Island by 1800—where the ‘funk’ but not the bird remains to this day. The last individual was killed in Iceland in 1844, leaving its current genus name Puinguinis as the final remnant of its life as the first penguin.

SOURCES

  • Cook R (1993) The Voyages of Jacques Cartier. Toronto: University of Toronto Press.
  • Gaskell J (2001) Who Killed the Great Auk. Oxford: Oxford University Press.

  • Grieve S (1885) The great auk, or garefowl (Alca impennis, Linn.): Its history, archaeology, and remains. London: TC Jack. [available here]

  • Thier K (2007) Of Picts and penguins—Celtic languages in the new edition of the Oxford English Dictionary. pp 246-259 in Tristram HLC (ed.) The Celtic Languages in Contact. Potsdam: Potsdam University Press

FOOTNOTES

  1. On Sat 19 Oct 2017, I gave a talk called ‘Discovering Birds in the Great White North‘ as part of a Bird Festival at the lovely Ruthven Park National Historic Site in the Niagara Region of Ontario. That talk drew material from a chapter I wrote on the History of Ornithology in Nunavut for a forthcoming book on the birds of Nunavut.
  2. This quote is from Cook 1993:xvii
  3. This quote is from Thier 2007:255

Gone Birds

BY: Bob Montgomerie, Queen’s University | 4 September 2017

martha_last_passenger_pigeon_1914
Martha in Cincinnati Zoo 1915

Last Friday, September 1st, was the anniversary of the death of Martha, the last Passenger Pigeon–a species that was, for centuries, the most abundant bird in North America. Martha was probably born in captivity in Charles Otis Whitman‘s aviary in about 1885, and died in the Cincinnati Zoo in 1914. I say probably because there is some debate about her origin (Schorger 1955). However, I think there are good reasons to consider Lord Rothschild’s (1907) account about her being raised by Whitman to be correct.

On the centenary of Martha’s passing, three excellent books (Avery 2014, Fuller 2014, Greenberg 2014) summarized just about everything you might want to know about Martha and the Passenger Pigeon. All of these books are worth reading for their different perspectives and the various ideas about why this most-abundant of species went extinct.

There can be little doubt that humans caused the extinction of this bird, but the process responsible for their million-fold decline, from billions in the 1870s to thousands by the 1880s, is still a bit of a mystery. Some have suggested that they were just wiped out by relentless hunting as were the Great Auk, the Labrador Duck, and the Dodo, for example. But those species were relatively rare (Labrador Duck and Dodo) or flightless (Dodo and Great Auk), unlike the Passenger Pigeon.

Others have wondered whether the pigeon’s demise may be an example of the Allee Effect whereby individual fitness declines with population size leading eventually to the extinction of local populations. While the Allee Effect might explain the extinction of some small colonies, it is hard to imagine how it would cause a species that still numbered in the thousands to go extinct across its entire range.

A new study by Ben Novak (2016) at UC Santa Cruz presents some intriguing new information that might help us to understand this massive extinction event. Genomic analysis reveals that the Passenger Pigeon probably numbered in the billions for at least 30,000 years, during a time when the forest ecosystems in its eastern North American breeding range were changing dramatically. Because the birds relied on masting tree seeds (oak acorns, chestnuts, pine seeds, maple seeds) as their main food supply, Novak suggests that the birds might have had a dramatic effect on forest ecosystems. By consuming a large portion of the plants’ reproductive outputs but also by depositing several inches of guano each year under their roosting and nesting sites, the pigeons may have been agents of habitat disturbance and destruction. For example, their massive roosts and nesting colonies broke trees and branches, opening up the canopy, and the several inches of guano they left on the ground, may have increased the incidence of forest fires. This sort of habitat disturbance may have been one reason they were so nomadic, returning to previous roosting and nesting sites only 5-20 years later, after the forests had recovered.

Passenger Pigeon populations were decimated by hunters, but Novak’s findings suggest to me that their final decline to extinction may have been due to both habitat destruction by humans settling eastern North America, and by the birds themselves. With much of their roosting and nesting habitat cleared for agriculture, the birds may have found their nomadic lifestyle simply unsustainable.

Clearly, the Passenger Pigeon still has stories to tell us about the causes of extinctions. Though this species is gone, we are fortunate to have specimens like Martha that can be mined for genomic data and the application of yet-to-be-invented tools. Some researchers have even suggested that we use these genomic tools to recreate and re-wild extinct species, but I am neither enthused nor optimistic about that prospect. Moreover, if Novak is correct, the forest ecosystems of eastern North America are now unlikely to be able to support a species that they last interacted with almost 150 years ago.


References

Avery M (2014) A Message from Martha: The Extinction of the Passenger Pigeon and Its Relevance Today. Bloomsbury, London.

Fuller E (2014) The Passenger Pigeon. Princeton University Press, Princeton, NJ.

Greenberg J (2014) A Feathered River Across the Sky: The Passenger Pigeon’s Flight to Extinction. Bloomsbury, London.

Novak BJ (2016) Deciphering The Ecological Impact Of The Passenger Pigeon: A Synthesis Of Paleogenetics, Paleoecology, Morphology, And Physiology. Mc thesis, University of California at Santa Cruz. Accessed 3 Sept 2017 from http://escholarship.org/uc/item/3260s35t

Rothschild W (1907) Extinct Birds. Hutchinson & Co., London.

Schorger AW (1955) The Passenger Pigeon: Its Natural History and Extinction. University of Wisconsin Press, Madison, WI.