Thursday, 26 June 2014

In Search of a Penguin’s Egg. Why?...Continued

Since writing In Search of a Penguin’s Egg. Why? on 8 April, I have read more and more on antarctic exploration in order: first, to understand why Edward Wilson, chief of the scientific staff of Scott’s final (1910-1912) expedition and who died with Scott, was willing to go to such lengths to obtain embryos of the Emperor Penguin; second, to try to understand why the Falkland Islands Dependency Survey (FIDS) was still trying to collect embryos in the late 1950s. I have also found, in the literature, the fate of the embryos collected by Nelson Norman in 1959.

However, the Scott expeditions (Discovery, 1901-04; Terra Nova, 1910-1912) were not the first to collect penguin embryos. Robert Neal Rudmose Brown (1879-1957) and James Hunter Harvey Pirie (1878-1965), members of the Scottish National Antarctic Expedition led by William Speirs Bruce (1867-1924), collected embryos at different stages of development of the Gentoo (Pygoscelis papua) and Adélie (P. adeliae) penguins. These embryos were brought back to Edinburgh where they were examined by David Waterston and Auckland Campbell Geddes of the anatomy department ‘through the kindness of Dr W.S. Bruce’.

The Scottish National Antarctic Expedition left Troon on 2 November 1902. Scotia, formerly the Norewegian whaler Hekla, had been refitted for the antarctic at the now defunct Ailsa shipyard in Troon, a short distance from where I shall be teeing off from the 1st at Royal Troon at 7.50, as usual on Fridays, tomorrow morning (weather permitting, of course). This successful expedition returned on 21 July 1904 to the Marine Biological Station at Millport, now sadly reduced to a study centre for schools. I shall return to the Scottish National Antarctic Expedition later since its very existence has a bearing on later developments.


Route of the Scotia

Scotia
Waterston and Geddes wrote up their findings for the expedition’s report published on 21 October 1909.

With regard to these developmental facts the question arises:— Is the duck's or the penguin's wing the more direct descendant of the common ancestor; or have they both diverged from the common stock approximately equally, but in opposite directions. Embryology alone cannot answer this question, but the evidence is clear in this, that the fore limb of the penguin in its development goes through a progressive and continuous series of stages along one unbroken line. The mesoblastic portion of the fore limb elongates, but its characters do not alter. It elongates, however, with a relatively greater rapidity towards the end of development, whereas the duck's fore limb, after being relatively longer than the penguin's ever is, regresses rapidly. So that the answer to our question, so far as the embryological evidence is concerned, must be that the wings of both these birds are different from the ancestral wing, and that the differentiation has been in opposite directions and that the common ancestor was a flying bird of a somewhat primitive type depending in large measure for the spread of its wing upon bone and muscle.
Their conclusions could not be more clear; the ancestor of modern birds and penguins was a flying bird.
One of the two laboratories on Scotia

Bruce had placed his penguin specimens with two of the leading anatomists/embryologists in Britain. David Waterston was demonstrator in anatomy at Edinburgh. In 1909, the year of publication of the report, he went to the chair at King’s College, London but in 1914 moved to the Bute chair of anatomy at St Andrews (as a golfer he perhaps could not resist the temptation of the Old Course). He died at the age of 70 in 1942, still occupying the Bute chair.

Coats Land. The expedition named this part of the
continent for the Coats family, the sewing cotton
manufacturers (now the multinational Coats plc),
who funded the expedition.
Auckland Campbell Geddes went on to a remarkable career. At the time of the penguin work he was assistant to the Professor (only one per department in those days), D.J. Cunningham, author of the famous Manual of Practical Anatomy. On graduation in 1908, he was appointed demonstrator. He was late to qualify because he had interrupted his medical studies to serve in the Highland Light Infantry during the Boer War.

His first public achievement from 1908 lived on until recently. His suggestion that the army volunteer force should be named The Territorial Force was adopted by Lord Haldane (JBS’s uncle), then Secretary of State for War. In 1909 he went to Dublin as Professor of Anatomy and then, in 1913, to McGill University in Montreal. He returned on the outbreak of war. He rose from Major in the 17th Northumberland Fusiliers in 1914 to Brigadier-General and Director of Recruiting at the War Office in 1916-17. He was then elected Member of Parliament for Basingstoke and Andover in 1917, becoming Minister of National Service from 1917 to 1919. He was then British Ambassador to Washington from 1920 to 1923. Sadly, he lost the sight of one eye. After his recovery, he served as Chairman of Rio Tinto. in 1942 he was injured by a flying bomb and also lost the sight of his other eye. He became Baron Geddes of Rolvenden in 1942. He died in 1954, aged 74.

The collectors of the penguin eggs also became well established. Rudmose Brown (1879-1957) was botanist on the Scotia Expedition. After working for Bruce after their return, he turned to geography, becoming Lecturer at Sheffield in 1907 and Reader at Manchester in 1920. He returned to Sheffield as Professor of Geography in 1931, retiring in 1945. In both World Wars, he served in Admiralty intelligence. Throughout, he continued his interest in the Arctic and Antarctic. He died in Sheffield in 1957.

Pirie (1878-1965) was the expedition’s medical officer, geologist and bacteriologist. On his return he was a medical practitioner in Edinburgh until 1913 when he joined the Colonial Service as a bacteriologist in Kenya. He was Deputy Director of the South African Institute for Medical Research from 1926 until 1941. In 1940 he gave the name Listeria (after Joseph Lister) to the bacterial genus. In 1948 he was living in Bournemouth on the south coast of England.

So, did Wilson not realise that the findings from the Scotia expedition would render his quest for Emperor penguin embryos unnecessary? Or did he judge the Emperor to be so different, in fact unrelated to the Adelie and Gentoo, that he thought the findings irrelevant? Was he even aware of the results from the Scotia expedition? Was he too busy with his work on the grouse disease inqjuiry and preparing for the Terra Nova expedition to study Waterston and Geddes’s findings between their publication in October 1909 and the departure of the Terra Nova in June 1910? I have been unable to find the answers to any of these questions from reading about Edward Wilson.

However, here’s where the politics of polar exploration kicks in. The Scotia expedition and its leader, Bruce, were regarded as the lowest forms of life by the power broker in British antarctic exploration, Sir Clement Markham, President of the Royal Geographical Society. All my reading indicates that Markham was rightly considered by later commentators a classic Monster. However, Markham’s personality is beside the point. The animus to Bruce was so great that I find it difficult to see Wilson raising any of the (excellent) results of that expedition in discussions over whether it was worthwhile to attempt the journey to Cape Crozier to collect the Emperor embryos with Markham in full control of the expedition and Scott, Wilson’s friend but Markham’s protégé.

From what I have read so far, I conclude that the collection of Emperor embryos was Wilson’s big ‘thing’ and had been his big ‘thing’ since his earlier discoveries on this species in the Discovery expedition of 1901-1904.

Geoge Seaver, in his book on Wilson, states that he gave three lectures to fellow expedition members and that ‘his own notes of them have been preserved but those taken by Scott and others are not without interest…In his second lecture he traced the descent of Penguins from the primitive lizard-bird, explaining their anatomy, and finding corroborative evidence in their primitive fossilized remains’. Those notes could well be worth examining anew to see what Wilson’s understanding was at that time.

Apsley Cherry-Garrard, Wilson’s protégé, who with Wilson and Bowers undertook that indescribable winter journey from Cape Evans to the rookery at Cape Crozier wrote in The Worst Journey in the World:
What is this venture? Why is the embryo of the Emperor penguin so important to Science? And why should three sane and common-sense explorers be sledging away on a winter's night to a Cape which has only been visited before in daylight, and then with very great difficulty?The Emperor is a bird which cannot fly, lives on fish and never steps on land even to breed. It lays its eggs on the bare ice during the winter and carries out the whole process of incubation on the sea-ice, resting the egg upon its feet, pressed closely to the lower abdomen. But it is because the Emperor is probably the most primitive bird in existence that the working out of his embryology is so important. The embryo shows remains of the development of an animal in former ages and former states; it recapitulates its former lives. The embryo of an Emperor may prove the missing link between birds and the reptiles from which birds have sprung*.
After the deaths of Scott, Wilson, Bowers, Oates and Edgar Evans on their return from the Pole, Cherry-Garrard obtained a series of embryos from Adélie Penguins at Cape Royde. He wrote:
It was always Wilson's idea that embryology was the next job of a vertebral zoologist down south. I have already explained that the penguin is an interesting link in the evolutionary chain, and the object of getting this embryo is to find out where the penguins come in. Whether or no they are more primitive than other nonflying birds, such as the apteryx, the ostrich, the rhea and the moa, which last is only just extinct, is an open question. But wingless birds are still hanging on to the promontories of the southern continents, where there is less rivalry than in the highly populated land areas of the north. It may be that penguins are descended from ancestors who lived in the northern hemisphere in a winged condition (even now you may sometimes see them try to fly), and that they have been driven towards the south. 
If penguins are primitive, it is rational to infer that the most primitive penguin is farthest south. These are the two Antarcticists, the Emperor and the Adélie. The latter appears to be the more numerous and successful of the two, and for this reason we are inclined to search among the Emperors as being among the most primitive penguins, if not the most primitive of birds now living: hence the Winter Journey. I was glad to get, in addition, this series of Adélie penguins' embryos, feeling somewhat like a giant who had wandered on to the wrong planet, and who was distinctly in the way of its true inhabitants.
So Cherry-Garrard in 1922 seems to be expressing a different reason for collecting embryos—that the most southern could be the most primitive penguins but not the most primitive bird—from Wilson in 1910. But the phrase if not the most primitive of birds now living could be taken to mean that it still might be shown that the Emperor is the most primitive bird. Alternatively, if not actually means even if not.  I tend to think it means the latter since otherwise he would be contradicting himself in these paragraphs.

Cherry-Garrard collected embryos from the Adélies during that period after the remaining members of the expedition found the bodies of the polar party, Scott, Wilson and Bowers. Surely Wilson in the many months they had together would have mentioned to Cherry-Gerrard whether Adéle embryos had been examined previously. That leads me to think that Wilson had not seen the report of the Scotia Expedition before embarking on Terra Nova. Original documents and notes on the contents of the libary taken by Scott and his party could well provide more information.

The story of penguin embryos does not end there…

----------------------
Waterston D, Geddes AC (1909) Report upon the anatomy and embryology of the penguins collected by the Scottish National Antarctic Expedition. Transactions of the Royal Society of Edinburgh 47 part 2, number 10, 223-244

Seaver, G (1933) Edward Wilson of the Antarctic. Naturalist and Friend. London: John Murray

*There is a difference in wording between the Pimlico edition 2003 (Vintage 2010) and that available online as a Project Gutenberg eBook; the latter appears to be from the original 1922 edition published by Constable in 1922.

‡Excellent photographs of the expedition are at: