Thursday 28 March 2019

Stephens Island: Lyall’s Wren—Scientific Pride and Prejudice in Victorian London. The Buller-Rothschild Feud

I had not done any homework for a circumnavigation of the south island of New Zealand. The reason I had not done so is simple. We were not meant to make that voyage but the storms in the Southern Ocean were so severe that a tour of the sub-antarctic islands had to be abandoned. As it was I did not notice passing a small island of historical significance to anybody interested in birds and their extinction. Had the sea been calmer in an area with infamously high winds and strong currents, the plan was to launch the Zodiacs and have a close look at the island but it wasn’t and afternoon tea (Devonshire cream tea plus sandwiches and cakes, since you ask) went uninterrupted.

The island is Stephens Island off the northernmost tip of the Marlborough Sounds and known to all reptile enthusiasts as one of the strongholds of the Tuatara. And no, I have never seen a Tuatara in the wild although a familiarity with its diapsid skull was an essential part of classical zoology; most university museums had a specimen. It is also famous as the final refuge and site of extinction of the flightless Lyall’s Wren (Traversia lyalli).

The story of the wren, the lighthouse keeper and the cats has been expanded by some urban myths but the whole story of how the last individuals of this small bird were first discovered in 1894, collected and then wiped out by feral cats only two years later is best described in this Wikipedia article which pulls together the facts and likeliest scenario. On mainland New Zealand the flightless wren had already been wiped out, probably by the Pacific Rat (Rattus exulans) which arrived along with the Maori from Polynesia in the 13th Century.

The scientific name itself is a reminder of the bitter feud it caused between two leading ornithologists of the day, Sir Walter Buller FRS and Walter, later Lord, Rothschild FRS. Naming new species was the name of the game in Victorian ornithology and the name, Traversia lyalli, was  given by Rothschild. The rule of the game is that the first to publish the new name takes the prize. If somebody else publishes a new name later, that name does not stand.

The animosity arose because Buller was shown the first specimen to be collected by the lighthouse keeper, David Lyall. Buller, who recognised it instantly as a new species, borrowed the dead bird  and sent it to the illustrator J.G. Keulemans in London. Buller then sent off a paper to the Ibis, complete with illustration, describing and naming the species as Xenicus insularis. Lyall then collected other specimens but sold them to the dealer, Henry Travers. Travers shrewdly reckoned that Lord Rothschild would be willing and able to pay more than Buller. He was right. Rothschild bought the nine birds and had them shipped to England. Rothschild then rapidly wrote a description of the bird—and named it Traversia lyalli—for a meeting of the British Ornithologists' Club. Ernst Hartert read the paper on behalf of, and in the absence of, Rothschild. Philip Lutley Sclater FRS was both the Club’s President and editor of the Ibis. He told Hartert that Buller’s paper for the Ibis was in the press but Hartert said he could not withdraw the paper without Rothchild’s consent. Well, he would say that, to quote Mandy Rice-Davies; Hartert was on Rothschild’s payroll.

After the Club’s meeting on 19 December 1894, the Bulletin containing Rothschild’s paper was published, astonishingly quickly, 10 days later, on 29 December. Buller’s paper, in the press since before the meeting of the Club, had to wait until April 1895 for publication in the Ibis.

Buller, not surprisingly, was described as incandescent with rage. He continued to call the bird Xenicus insularis for the rest of his life and attacked Rothschild, with whom he had dealt with—and sold bird specimens to—at every opportunity.

Keulemans's illustration for Buller's paper in the Ibis

Keulemans depicted a male and female for Buller's last book -
the Supplement in 1905-06

Looking at Rothschild’s behaviour nearly 125 years after the event, I would have to ask. Was it possible for Hartert to contact Rothschild after the meeting and before publication to tell him what had happened? If so, did he do so? And would Rothschild have had time to withdraw the paper or to change the wording to acknowledge Buller’s real priority and name for the bird? If the answer to the last question is ‘Yes’, then I have Lord Rothschild down for conduct unbefitting a gentleman.

Walter Rothschild
Buller versus Rothschild continued on this and other matters ornithological, with every opportunity being taken in print to have a go at the other. The best analogy I can think of is a Twitter War in slow motion.

The vituperation continued after Buller’s death in 1906. Rothschild’s book, Extinct Birds (Hutchinson: London) was published in 1907. Under literature referring to extinct birds he wrote:

1905-1906. Sir Walter Buller. Supplement to the “Birds of New Zealand."Two volumes. (Though containing very interesting notes on extinct and threatened birds, these two volumes are rather disappointing. They contain very little that is new, and are mainly composed of quotations from other people's writings or letters. Buller's former great book on the Birds of New Zealand was a most important and creditable work, though not without shortcomings. Our knowledge of New Zealand Birds might have been brought up to date in his supplement, but we cannot say that this has been done properly, and errors are frequent.) 

And on the still extant birds, the Chatham Island Robin (Petroica traversi) and Snares Tomtit (P. dannefaerdi):

Sir Walter Buller…has confounded M. traversi and dannefaerdi, and the figure he gave on his plate looks so black, that I do not doubt it represents rather the latter than the former. Of course M. dannefaerdi alone occurs on the Snares, and Buller's traversi from the Snares were all dannefaerdi. Dr. Finsch's statement (Ibis 1888, p. 308) that Reischek's specimen from the Snares "agreed in every respect with specimens from the Chatham Islands" is entirely wrong, for, even if one prefers unscientifically to lump allied forms, one cannot say that a Miro from the Chathams agrees in every respect with one from the Snares. Buller's doubts about the distinctness of the latter might easily have been removed, if he had taken the trouble to compare them, for it does not require any genius to see the differences. I admit that with my present views on geographical forms I would regard the two Miro as sub-species, and call them M. traversi traversi and M. traversi dannefaerdi, but most ornithologists would still consider them to be "good species."  
I may add that Buller…has not quoted my description correctly, for in his rendering are several disturbing misprints, and in the fourth line from the bottom occurs a "not" which ought not to be there, and which makes the sentence incomprehensible. Also the name itself is spelt incorrectly. 

Rothschild’s take on the by-then-extinct Lyall’s Wren story:

I received nine specimens of this new bird, and was not aware that any others had been taken at that time. As I was unable to attend the December meeting, 1894, of the British Ornithologists Club, I asked Dr. Hartert to exhibit the birds in my name. When he had done so and had read the description, the Chairman, Dr. P. L. Sclater, said that the bird had also been received for illustration and description in the Ibis, from Sir Walter Buller, and he asked Dr. Hartert if I would not withdraw my description. Dr. Hartert said that this was unfortunate, but he had no authority to withdraw my description, and he and Dr. Sharpe thought that the proceedings of the meeting should be printed without consideration of any manuscripts which might refer to the same bird. No doubt this was hard luck on Sir Walter Buller, but it would have been equally hard luck for me if he had forestalled me with the new bird. He had only one specimen, I had nine, of both sexes, and I had paid a high price for them, as types of a new bird. My type is in Tring, and, as everybody knows, available for study by any competent ornithologist, while Buller's type was not in any museum, and it was uncertain to whom he would sell it afterwards. I suppose it is now in the Carnegie Museum, Pittsburgh, to which Buller's "third collection," 625 specimens, was sold for a thousand pounds, as Buller himself tells us in his Supplement II, p. 167, under the heading of Glaucopis wilsoni!

On the same page Sir Walter Buller also tells us that his "second collection" was sold to me, but he makes a mistake about the price, as I certainly did not pay a thousand pounds for it. 
I mentioned these unimportant details, because Buller rather bitterly and severely complained about my describing the Stephens' Island Wren, on p. 111 of his supplement. I may only add that of course my name, being published in December, 1894, has the priority over his, which was not published before April, 1895. 

Another well-aired disagreement was whether there were two species (one North Island, one South) of the now extinct Laughing Owl (Sceloglaux albifacies):

Having thus discussed the age of this owl, the question must be considered if it is different from S. albifacies from the South Island. This is less easily done. Buller described it as a "new species," and mentions among the distinctive characters (see above) the colour of the tail. The tail, however, is "skillfully" (as Buller calls it, though I should use a less complimentary adverb) stuck in, and does not belong to a Sceloglaux, but to an Australian Ninox, and also some feathers on the neck are foreign.

There are other examples but that’s enough for anybody. The whole sorry story, the owl, the wren and Buller's sales and attempted sales of specimens to Rothschild (the latter had a flock of live kiwis supplied by the former), can be found in Errol Fuller's book, Extinct Birds

Keulemans also illustrated Rothschild's book. A pair of
Lyall's Wrens in the corner of a plate

So while Rothschild thought he was entitled to name the bird because he ‘had paid a high price for them’ we can only marvel at his sense of entitlement. The wren’s recently-studied DNA does though indicate that Rothschild was right to have erected the genus Traversia, separate from Xenicus.

Unfortunately, none of the acrimony was of any use to the bird itself. It was dead; an ex-wren; extinct, lingering only as dead specimens in museums to remind us of the great damage done to the original fauna of New Zealand.

Fuller E. 1987. Extinct Birds. London: Viking/Rainbird.

Galbreath R, Brown D. 2004. The tale of the lighthouse-keeper’s cat: Discovery and extinction of the Stephens Island wren (Traversia lyalli). Notornis 51, 193-200.

Updated 9 April 2019

Tuesday 26 March 2019

Buller’s Birds: Spotted in New Zealand

Buller’s Albatross—Tick. Buller’s Shearwater—Tick. Buller’s or Black-billed Gull—Tick.

In, and off the coast of, New Zealand it is easy to see the full set of birds named after Buller. I really had not thought about Buller until we saw these species over a period of a few days earlier this year.

Sir Walter Buller
Portrait by Ethel Mortlock
Sir Walter Buller (1838-1906) was a social climbing lawyer, government administrator and amateur ornithologist who, I was surprised to discover, was elected to the Royal Society in 1879. He seems to enjoy a mixed reputation in New Zealand, mainly, it has to be said, not good.

Ornithology is one of those ‘ologies’ that does not mean what it says. Thus, ornithology has meant—and continues to mean—not the study of all aspects of the life of birds but what those who called themselves ornithologists defined ornithology to be. Some areas of biology are ‘in’; others are ‘out’. At the time of Buller ornithology was the collection of birds for museums (public and private institutions), their distribution and their taxonomy. The only part of the bird (apart from eggs) that was preserved was the skin. Because there was great rivalry between institutions and wealthy individuals for specimens, there was an active market. Collectors in exotic places shot and trapped large numbers of birds—the rarer the better—to sell to the museums.

Buller became the leading expert on the ornithology of New Zealand.

The son of Methodist missionaries, he started as a a native interpreter in the magistrates’ courts. but by self promotion and ability he rose in the department of government dealing with the purchase of Maori lands. Throughout this time he worked on his birds to the extent that he had enough material to write a major book. In 1871 he obtained a government grant, leave on half pay and a job with the New Zealand agent in order to go to London to arrange publication.

A website on New Zealand history has the story:

Buller drew up an initial outline for his book in July 1867. At about the same time his ornithological work was subjected to its first critical analysis. A German ornithologist, Otto Finsch, had obtained a copy of an essay Buller had distributed to various institutions in London. Buller had prepared the essay for the New Zealand Exhibition in 1865 but this was the first time it had been made available in print. Finsch, who had 'made a special study of the birds of the Pacific Islands', took issue with some of the new species Buller identified. His attempt to analyse and classify species was at odds with the approach of most contemporary New Zealand scientists, who were prepared to leave this responsibility to the ‘experts' in Europe. Buller fought off the criticism but also realised that for his book to be taken seriously he would have to base himself in London and make use of the superior library and museum facilities there. 
Buller didn't revive his book outline until July 1870. By this time Dr James Hector, then 'the key figure in New Zealand science', was less enthusiastic about a proposal he and Buller had worked out in 1867. This entailed Buller donating his specimens (then numbering over 200) to the Colonial Museum, in exchange for the government contributing £300 towards the book's publication. Hector was particularly concerned that Buller's specimens were worth less than they had discussed. Cabinet accepted Hector's recommendation that Buller be required to provide the government with 25 copies of his book and approved the grant. The following year Buller asked Dr Isaac Featherston, who had seconded him to work on the Rangitikei-Manawatu purchase, to support his application for special leave from his position as resident magistrate in Whanganui. Buller was granted 18 months leave on half pay plus a return passage to London, where he arrived in October 1871. 
As part of the agreement Buller also negotiated a position as part-time private secretary to Featherston, who was due to be appointed New Zealand's new agent general in London. His salary was to be equivalent to his total income at Whanganui. This was fuel for the Opposition, who accused the government of patronage. They seized on the fact that the new magistrate at Whanganui was receiving only a third of the £600 Buller was getting while on leave in London, and noted that Buller was earning more in London that he had in Whanganui. Though Buller explained the disparity as a mistake in his estimate of court fees, he was eventually forced to resign his magistrate's position. Featherston was instructed not to employ him as secretary. The Evening Post scornfully reported on the situation on 18 August 1873: 
Mr Walter Buller has hitherto been one of Fortune's favourites in the Civil Service – one of those curled darlings who must be provided for at all hazards, and who had only to open his mouth in order to have it filled with plums.

However, when his book, A History of the Birds of New Zealand, was published in parts in 1872-73, it was so highly praised that the British Government awarded him a civil honour, Companionship of the Order of St Michael and St George, in 1875. His more senior and unrewarded colleagues back home in New Zealand were left, apparently, spitting feathers.

The book was also a commercial success. He sent the 25 copies promised to the New Zealand administration with a note pointing out that all but 40 of the 500 printed had been subscribed for (subscribers included Charles Darwin). But London had more to offer than a publisher for his book; he read law and was called to the Bar in 1874.

When Buller returned to New Zealand he practised as a barrister. By all accounts his advocacy in the Native Land Courts was controversial but highly lucrative, so lucrative that he retired from the law and moved back to London to become a  commissioner for the Colonial and Indian Exhibition of 1886. For that his CMG was raised to KCMG. He became Sir Walter Buller.

Throughout his professional life, his work on birds continued apace. A new, enlarged, edition of his book was published in 1888.

After what are described as business difficulties took him back to New Zealand in 1890 he became embroiled in a legal dispute over land he had bought from the Maori Te Keepa. In defending and winning what actually his own case he acted for Te Keepa—and charged him for the privilege.

Buller then returned to England in 1899 enjoying the life of a gentleman naturalist in the social scene of late Victorian London.

Apart from what are now regarded as dodgy dealings over Maori land deals his counsel of despair over the fate of New Zealand’s native wildlife and his dealing in birds is perhaps what he is best remembered for.

Buller was of the opinion that everything native to New Zealand would be replaced by the more vigorous European immigrants. Introduced British birds and mammals were spreading through the newly-created fields and non-native forests. The Maori, he argued, were doomed. He therefore opposed conservation of both native forests and birds as a hopeless exercise. Eventually he did back pleas for the statutory protection of native birds and the creation of sanctuaries on two offshore islands. Although he backed these efforts, Buller remained unconvinced that they would have any significant effect and continued to collect specimens of the rarer birds for his own studies (‘in the interests of science’) and others. Selling skins to collectors throughout the world was highly lucrative.

Buller’s spectacular falling out with Lord Rothschild, that great collector with a private museum, to whom he sold skins, and their bickering in print, I will write about later.

Buller’s links with Otto Frisch (1839-1917) in Germany are noted above, with Frisch objecting to Buller’s erection of some new species. However, there must have been some rapport because Buller was awarded, with Finsch's assistance, a doctorate in natural history by Tübingen University in 1871. Finsch, though, like Buller, had political interests as well as professional ornithological ones. He was involved in German colonial expansion in the Pacific and as Bismarck's Imperial Commissioner in 1884 he negotiated the deal whereby the north-east of New Guinea together with some islands became a German protectorate.

Even a quick perusal of his major work (first edition online here) shows the immense amount of work that went into it, all of it gathered while employed in the civil service. After the second edition (1887-88), he produced a two-volume Supplement in 1905 to bring it up to date.

I am by no means convinced that Buller deserved the brickbats he has received in New Zealand in recent years. He was, after all, an ornithologist—a highly gifted and dedicated ornithologist—doing what ornithologists of the time did. Yes, he was a pessimist, arguing that the extinction of native species—which did occur—was inevitable. But anybody visiting New Zealand today can see the reason for that pessimism: a mainly artificial landscape devastated by the spread of agriculture and non-native forestry; two waves of human migration both of which resulted in the killing of spectacular native birds for ceremonial purposes or museums; the introduction of alien species some of which like the Stoat, Weasel, Brush-tailed Possum, mice and rats plus the domestic cat kill the native birds. New Zealand is trying very hard, starting with the predator-cleared islands, to prevent further extinctions and to promote the restoration of habitat throughout the country. So far it has done a brilliant job but the ambition to rid the entire country of its alien predators has a very long way to go.

And here are Buller's eponymous species:

Buller's or Black-billed Gull (Larus bulleri) - left
From Buller's book. Illustration by John Gerrard Keulemans
Buller's Albatross (Thalassarche bulleri)
Taken by a member of the Noble Caledonia Expedition Team
on board MV Caledonian Sky 28 January 2019
Buller's Shearwater (Ardenna bulleri)
Photo Aviceda at English Wikipedia

Friday 22 March 2019

Galapagos Iguana Hybrids: What about their Salt Glands?

Last week I gave a talk on salt glands to a local group. I remembered that I had not raised a question here that arises from the existence of some very odd animals on one island of the Galapagos.

Everybody knows there are the Marine Iguanas (Amblyrhynchus cristatus) and Land Iguanas (Conolophus sp.) on the Galapagos Islands. Evidence indicates they had a common ancestor about 4.5 million years ago. Watchers of television wildlife programmes cannot fail to have noticed the entirely different lifestyles. Land iguana eat plants and do not venture into the sea. By contrast, Marine Iguanas eat seaweed often foraged at considerable depths.

Over the past forty years it became evident that on one very small island, Plaza Sur, just off the east coast of Santa Cruz (Indefatigable) there were hybrids present between Marine and Land Iguanas (Conolophus subcristatus). These hybrids are sterile as one might expect. There is good evidence that their fathers are male Marine Iguanas, their mothers Land Iguanas. It seems that the breeding seasons of the two species can overlap there. The habitats are very not very far apart either, the thin vegetation where the Land Iguanas bask and feed being only a matter of yards from the rocks on which the Marines bask.

We took video and still photographs of one of these hybrids on Plaza Sur in January 1012. It was lying with Land Iguanas. Local guides say they are not very active and typically have a ‘spaced out’ look like the one below:

Hybrid Marine x Land Iguana, Plaza Sur, 12 January 2012

For comparison: Galapagos Land Iguana, Plaza Sur, 12 January 2012

Also for comparison: Galapagos Marine Iguana, Plaza Sur, 12 January 2012

Nesting burrows of Land Iguanas. Plaza Sur, 12 January 2012

Vegetation on Plaza Sur, 12 January 2012. The marine iguana-like claws of the hybrid iguanas are said to be advantageous in reaching the fruits of the prickly pear. The land iguanas have to wait for them to drop

The question I have had since then is what do the salt glands of these hybrids do? Marine Iguanas have very active salt glands which operate to remove the sodium chloride ingested during feeding. In addition, the concentration of potassium is about ten times higher than in birds which could reflect the high potassium concentration in the plants of their diet. Marine iguanas on land can be seen snorting secretion from the salt glands out of their nostrils, a phenomenon observed by Charles Darwin who did not appreciate its significance to their survival.

Other, species of iguanid lizard have salt glands that can secrete both sodium and potassium. There is evidence that the ratio can be changed by varying the diet. Lizards on a high sodium diet or given a salt load produced a secretion with a more sodium than potassium. As far as I am aware, the Land Iguanas of the Galapagos have not been studied in this respect.

The hybrids seem to live and eat like their Land mothers although I see there are anecdotal reports of their being seen eating seaweed but not entering the water to do so.

So do the hybrid iguanas on Plaza Sur have functional salt glands? How does their size compare with the Marine and the Land? If the salt gland is functional, what are the stimuli for secretion, and does the ratio of sodium:potassium change to reflect changes in the dietary intake?

Possibilities of studying the hybrids to answer these questions are low but the insights into which genes are involved in salt-gland structure and function would be fascinating outcome of this ‘natural experiment’ but biological dead end.

I  used this drawing from Bill Dunson's paper in our book on salt glands

Thursday 21 March 2019

Protein deficiency in early life. Experiments from 1960s Hong Kong

When I was writing the post of 8 March on the classic experiments in malnutrition and the intergenerational effects that ensued, I could not help thinking I was vaguely aware of earlier experiments that also explored the effects of a shortage of protein in the diet on a rat’s ability to navigate. I then realised who was doing it and where but I had no idea of the background or the outcome. It was actually in the next room of the animal house we occupied at the University of Hong Kong (it was air conditioned!) and just getting started on it by the time we left was the new, and first, Professor of Psychology. He was known then as John Dawson. I only spoke to him a couple of times but I had him down as a good guy since he spoke in terms of evolutionary psychology. But what was he doing and why?

I knew for that he was using the colony of rats in the Department of Zoology, an inbred strain founded from animals flown in from the University of Sheffield about eighteen months earlier*. I also knew that the zoology workshop (brilliant with perspex constructions) had built the test maze.

John Binnie-Dawson in 1969
from a family tree on
From the university newsletters that arrived over the years (long discontinued) I realised that the John Dawson we knew was being referred to as Professor J.L.M. Binnie-Dawson. Therefore, in tracking what he did I had to use the two names since I discovered he published under both.

It is clear that he had observed the feminising effect of kwashiorkor, the result of extreme protein deficiency after weaning, on children in West Africa. Dawson had worked in Sierra Leone. From his publications it is also clear that he was interested in differences between the sexes in spatial ability and so the two fields were brought together in studying effect of protein deficiency and sex hormones in rats.

I have not read the papers, having been content to see if my memory was correct but it seems that he pulled the whole story together in a paper in Psychologia in 1984 of which this is the published summary:

The author presents empirical evidence, mainly from his own research conducted from 1963 to 1975, to support the biosocial hypothesis, which argues that the normal sex difference between males and females in spatial ability has a 2-way sex hormone basis. This higher level of male spatial ability is also held to be biologically adaptive for the human group: In subsistence hunting societies, the male must generally be able to hunt food and locate water to ensure the survival of the group. This hypothesis also postulates that the actual level of male spatial ability in a particular biological subsistence environment will be influenced by the spatial requirements of that environment. Three specific studies are reviewed: (1) a study to determine whether Arunta individuals living in the semi-desert environment of Central Australia have developed high spatial ability and a permissive social system, as contrasted with the strict group orientation of West African Temne rice agriculturalists, who do not need spatial ability to grow and accumulate rice; (2) a study of the effects of the Kwashiorkor-induced endocrine dysfunction and feminization of males on the development of feminine spatial ability; and (3) a study using rats to test this suggested endocrine basis of spatial ability under controlled laboratory conditions. It is concluded that these biosocial studies have provided extensive evidence for the biosocial hypothesis.

Other research interest included handedness, as well as a number of other areas in cross-cultural psychology, much of relevance to the people of Hong Kong.

While in Hong Kong, he was instrumental in proposing and then hosting in 1972 the inaugural meeting of what became the International Association fo Cross-Cultural Psychology. He was its first Secretary-General and later its President.

John Lewis Mervyn Binnie was born in 1930 in Sydney. According to genealogy websites the name Dawson was that of his paternal step-grandfather, Edwin Dawson, who adopted him after his father died and shortly before his mother died. This is the name he published under until 1980 when he changed his name to Binnie-Dawson.

Before he arrived in Hong Kong, he published from Edinburgh in the early 1960s where he was Research Lecturer in the Department of Social Anthropology; it seems it was while based there that he worked in Sierra Leone. He also served as a Lieutenant in the Royal Navy Reserve. Later in the 1960s, he was in the Department of Psychology at the University of Sydney. It was from there that he was appointed to the new chair in Hong Kong.

John Binnie-Dawson retired in the mid-1980s to Kingswear in Devon. He died in Torbay Hospital on 29 December 1987, aged 57.

John Binnie-Dawson was one of a small group who during the 1960s lifted the University of Hong Kong from being a largely teaching institution to one recognised internationally in research and scholarship.

…And he was another to demonstrate how important it is to be given a good start in life.

*The rats provided a good source of income for the department under a wheeze devised by John Phillips (1933-1987) (FRS 1981). Excess were preserved in large numbers and charged—in advance—to the ‘A’ level Examinations Board (then run by the University) against their possible use in practical examinations. The charge per rat was not small.

Binnie-Dawson, J. L. [1984]. Bio-Social and Endocrine Bases of Spatial Ability. Psychologia: An International Journal of Psychology in the Orient 27: 129-151.