Lemming Population Cycles 100 years after Charles Elton

2024, the centenary of the publication of Charles Elton’s classic paper, Periodic fluctuations in the numbers of animals: their causes and effects, in Journal of Experimental Biology, also saw a flurry of interest on the same topic and the same mammals that occupied a large part of Elton’s review, lemmings.

Norway Lemming, Lemmus lemmus
Kevo Strict Nature Reserve, Finland, 2011
by Lakahillo (from Wikipedia)

The spectacular periodic fluctuations in lemming numbers—each cycle lasting 3-4 years—described by Elton from evidence gathered from around the Arctic have excited renewed interest because there had been suggestions that the large cyclical fluctuations in abundance are being wiped out by recent climate change, with consequences for the populations of predators and the whole ‘ecosystem’. However, twenty-four authors from Canada, France, Sweden, Finland, Norway, Russia, USA, Germany and Denmark analysed 24 sets of recent data gathered across the Arctic that recorded population changes over time.

This is the authors’ summary:

Reports of fading vole and lemming population cycles and persisting low populations in some parts of the Arctic have raised concerns about the spread of these fundamental changes to tundra food web dynamics. By compiling 24 unique time series of lemming population fluctuations across the circumpolar region, we show that virtually all populations displayed alternating periods of cyclic/non-cyclic fluctuations over the past four decades. Cyclic patterns were detected 55% of the time (n = 649 years pooled across sites) with a median periodicity of 3.7 years, and non-cyclic periods were not more frequent in recent years. Overall, there was an indication for a negative effect of warm spells occurring during the snow onset period of the preceding year on lemming abundance. However, winter duration or early winter climatic conditions did not differ on average between cyclic and non-cyclic periods. Analysis of the time series shows that there is presently no Arctic-wide collapse of lemming cycles, even though cycles have been sporadic at most sites during the last decades. Although non-stationary dynamics appears a common feature of lemming populations also in the past, continued warming in early winter may decrease the frequency of periodic irruptions with negative consequences for tundra ecosystems.

Two commentaries were published on the paper by Charles Krebs and by Rudy Boonstra, well-known ecologists who work on population changes. These commentaries highlight the fact that despite decades of research the underlying cause of these population cycles in lemmings and other rodents remains unknown. Similarly, the physiological processes involved are also unknown. I have written previously on Dennis Chitty’s (1912-2010) attempts to identify the cause of population cycles in voles (see here).

What is clear is that the mathematical modelling approach pioneered by Alfred Lotka in 1925 based on changes in numbers of predators and prey over time and which have been pursued actively ever since are, as is so often the case with mathematical models, inadequate. Experimental approaches to lemming and vole cycles have also not, in general, been successful, with other possible explanations of any changes observed and doubt whether the experimental conditions imposed are ones that actually obtain in the wild.

The importance of trying to understand what happens under the snow—the habitat of lemmings in winter where they feed and breed—was also stressed in the commentaries. Those observing lemmings only see the results of that activity when they count the population in spring and summer.

It is easy to forget when discussing the cause of cyclicity in population size and the changes in reproductive rate necessary to drive such changes that the the occurrence of ‘lemming years’ is a major wildlife phenomenon. Elton wrote of the Norway Lemming, Lemmus lemmus:

For many years the lemmings have periodically forced themselves upon public attention in Southern Norway by migrating down in swarms into the lowland in autumn, and in many cases marching with great speed and determination into the sea, in attempting to swim across which they perish. The details of the fate of the migrants do not concern us here and are fully described by [R] Collett [1911-1912]….Lemming-years in Norway have the status of great floods or terrible winters.

and of lemmings in northern Scandinavia:

It is obvious that the phenomenon of migration is far more striking than a mere increase in the numbers. The spectacle of processions of lemmings ecstatically throwing themselves over the ends of railway bridges, and falling to an apparently useless death below; the sea strewn with dead lemmings like leaves on the ground after a storm; lemmings making a bee-line across crowded traffic oblivious to danger; all these things are bound to make people talk.

Is was descriptions like this that gave rise to the popular notion generated and churned by the news media that lemmings commit suicide rather than simply die en masse when presumably moving in large numbers in search of food (Elton wrote not of food but of ‘relieving congestion in an area’). The Walt Disney film White Wilderness (1958) showed the sort of scenes described by Elton and others, with lemmings tumbling over the edge of a cliff. Decades after I saw the film at a meeting of the school natural history society in 1960 it emerged that captured lemmings had been pushed over the edge of artificial rockwork. The footage may have been fake and the lemmings poorly treated but film certainly conveyed what happens in the wild.

There are questions in my mind on this related phenomenon of migration, like the distances involved and whether they are acting like a plague of locusts, stripping one area of food before having to move on, given that their body size and increased activity would suggest they would  starve to death within a week without food. However, it is the response to  overcrowding view that dominated research in this area for decades. But that is another story.

To conclude, 100 years after Elton, the cause(s) and mechanism(s) of population cycles in lemmings—and voles etc—remain unsolved. But research activity and monitoring continues. We can only hope for progress.

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As an afternote I found reading Elton’s paper from 1924 well worth while. Not only did he consider mechanisms that may account for the phenomenon in lemmings but he also reviewed the occurrence of periodic booms in population of other rodents in some years. In the garden we had a large population of Bank Voles in the summers of 2018 and 2019, but not in the years before or since. Are the same or different mechanisms at play in different species is another abiding question.

Boonstra R. 2024. Population regulation and limitation—insights from lemming cycles: past, present and future. Proceedings of the Royal Society B 291: 20240660. doi.org/10.1098/rspb.2024.0660

Elton CS. 1924. Periodic fluctuations in the numbers of animals: their causes and effects. Journal of Experimental Biology 2, 119–163. doi.org/10.1242/jeb.2.1.119

Gauthier G, Ehrich D, Belke-Brea M, Domine F, Alisauskas R, Clark K, Ecke F, Eide NE, Framstad E, Frandsen J, Gilg O, Henttonen H, Hörnfeldt B, Kataev GD, Menyushina IE, Oksanen L, Oksanen T, Olofsson J, Samelius G, Sittler B, Smith PA, Sokolov AA, Sokolova NA, Schmidt NM. 2024. Taking the beat of the Arctic: are lemming population cycles changing due to winter climate? Proceedings of the Royal Society B 291: 20232361. doi.org/10.1098/rspb.2023.2361

Krebs CJ. 2024. Lemming population fluctuations around the Arctic. Proceedings of the Royal Society B 291: 20240399. doi: 10.1098/rspb.2024.0399

Central American Agouti. Honduras 2024

 

We saw the Central American Agouti (Dasyprocta punctata) several times while walking in the gardens of the Lodge at Pico Bonito in Honduras. These hystricomorph rodents are diurnal and live as mongamous pairs. Most times we saw one it was carrying one of these large seed pods.

Commander Hughes, bird artist, and his cousin, the ornithologist Pat Hall

After my article on the artist Commander Alfred Marcus Hughes appeared, Greg Davies commented:

Hughes also makes a cameo appearance in Beryl Patricia Hall's autobiography "A Hawk from a Handsaw" (1993). Hall was the doyenne of the British Museum Bird Room in the post-war III period, and a relative of Hughes.

On looking up Hughes and Hall I found that they were first cousins, once removed. Pat Hall was the grand-daughter of Hughes’s mother’s brother.

The comment also rang a bell in my neural circuitry. I was sure I could remember meeting Pat Hall but could not recall where and how. Then, looking her up to write this article, I found a photograph which provided all the answers. The date was 4 May 1972 when she received the Stamford Raffles Award at the AGM of the Zoological Society of London. I remember speaking to her briefly after the meeting when she seemed a bundle of nerves. For some strange reason on seeing the photograph I immediately remembered the coat she was wearing as being several sizes too big such that she and the coat seemed moved independently—a very weird thing to recall from the memory bank after 52 years.

Pat Hall receiving the Stamford Raffles Award from the President of the
Zoological Society of London, Prince Philip, Duke of Edinburgh.
4 May 1972
(Annual Report for 1972, ZSL)

Pat Hall was Beryl Patricia Hall (née Woodhouse). She was born to a wealthy family in Epson, Surrey, in 1917. She was thwarted by her parents in going to Cambridge to study maths and instead spent four years at home kicking her heels but also developing an interest in birdwatching. Thus in the 1939 Register, the emergency census, she is shown as employed on ‘domestic duties’ but also as involved in Air Raid Precautions work teaching ambulance drivers. She was determined to do something for the war effort and joined first the Womens' Legion and then the Mechanised Transport Corps, a uniformed women’s civilian organisation that provided drivers to government departments and other civilian services. 

After the outbreak of war Pat Hall became engaged to John Clavell Hall, in civilian life a Winchester-educated insurance clerk. He was commissioned into the Royal Artillery in March 1940 and posted to the Middle East. Pat volunteered to serve overseas driving ambulances. She was first in South Africa but was then moved to Egypt. The two were married on 21 May 1941 at Suez. However, this was no happily-ever-after story. By June 1943 John Hall had fought in Crete. was a temporary Captain and had been mentioned in despatches. However later that year he was captured during the Battle of Leros in the Dodecanese Campaign. Pat further developed her interest in birds in North Africa and ended the war in Italy still driving ambulances. The MTC had been incororated into the Army during her time in Egypt and she ended the war as Sergeant-Major, with a  'Mention in Despatches', of 502 Motor Ambulance Convoy, skilled in driving, servicing and maintaining motor vehicles.  Her army life ended with the release of her husband in May 1945. However, the marriage broke down and it was then that Hughes must have offered his help.

Alfred Hughes was a friend of the Norman Boyd Kinnear of the Natural History Museum. Kinnear, with no academic qualifications, had risen through the ranks of the museum world, beginning at the Bombay Natural History Society’s museum in Calcutta and continuing at the Natural History Museum in London. He had specialised in ornithology. By the time the war was ending he had been appointed Keeper of Zoology. In 1947 he was asked to take over as Director, past the normal retirement age, until a suitable candidate had been found in the post-war world. He stayed on as Director until 1950, sorting out the repair and restoration of the Museum needed after the disruption and damage caused by the War; for this he received a ‘K’.

Through that contact with Kinnear, Pat Hall was offered a post of Associate Scientific Worker in the Bird Room of the Museum. Ideally suited for those wealthy enough not needing to find paid employment, the scheme provided cheap and enthusiastic voluntary labour to the Museum. The position was paid at a rate—which did not change for ten years—of four shillings per hour, a wage which would have exceeded the average industrial pay in the 1940s. However, instead of that equating to a salary of around £800 per year, the maximum pay each year was capped at £100. Pat Hall was clearly keen to work at the Museum; she turned down a job offer from the BBC in order to work at the Museum.

James David Macdonald (1908-2002) was in charge of the Bird Room. He recognised her aptitude for the work she was doing and given her experience in Africa and her skills as a motor mechanic invited her, at her own expense, to be a member of a collecting expedition to south-west Africa that began in late 1949. The team, including Macdonald’s wife as the team doctor and cook, collected 1300 specimens of nearly 200 species during the six months in the field. After that for nearly 20 years Pat Hall divided her time between curatorial work, collecting expeditions and research based on the collections in the Museum. Her expeditions during the 1950s were to Africa. In 1953 she organised and largely funded a Natural History Museum-backed trip to what is now Botswana. Before and after an ornithological congress in Livingstone in what is now Zambia, she collected in Botswana and Angola respectively; overall she collected around 2000 specimens for the Museum. A short trip to north-western Botswana was her final experience of collecting in Africa.

Her publications during the 1950s and 60s were on African birds. Her largest was the outcome of eight years of work: An Atlas of Speciation in African Passerine Birds in 1970. Of 423 pages with 439 maps it was widely praised by reviewers. Herbert Friedmann (1900-1987) the American ornithologist who worked at the Smithsonian Institution wrote of its importance in The Auk of July 1971:

As Mrs. Hall states at the beginning of her introduction, this great series of maps constitutes the first attempt to show in graphic form, for the ready comprehension of the student, the results, "and the continuing process, of evolution in a large continental avifauna by means of plotting on one map the distribution of species believed to be immediately descended from a common ancestor." By placing closely related species on the same map it becomes evident where they overlap and where the are allopatric, and these facts give the field student indications of where to look and what to study in an attempt to assess and to interpret the past history of each of these current distributional patterns. It must be kept in mind, when using this atlas, that every existing specific distributional picture is not merely a discrete fact of local interest, but is always, and inevitably, the result of the past history of the species and of the region. This is the real, inherent interest in each of them, and each could become a valid point of departure for further study of the evolutionary vicissitudes of the particular species.

The Atlas had started as a joint effort with Reginald Ernest Moreau (1897–1970) the great amateur scientific ornithologist of his day. However, he became increasingly ill and the work fell on Mrs Hall. He died in the year of its publication.

Work on African birds was interrupted in the early 1960s. Macdonald heard of and then approached Harold Hall (no relation to Pat) an Australian philanthropist who was interested in supporting research. Harold Hall agreed to sponsor a series of five expeditions for the Natural History Museum (still labouring under the title of British Museum (Natural History)) around Australia. Macdonald was very keen to obtain more specimens for the Museum. The number of Australian specimens available within Britain had been depleted by the sale of Lord Rothschild’s collection (Rothschild was being blackmailed by a ‘titled lady’ and he needed the money) and another private collection to the USA.

Pat Hall led the third Harold Hall Expedition given her experience of collecting in arid parts of Africa. That was to the interior of Western and South Australia. She then edited the book describing the findings of all five expeditions which was published in 1974.

After the post-war recovery the Natural History Museum on its site in London was bursting at the seams with many thousands of specimens stored in unsuitable accommodation. In the mid-1960s, plans emerged to move the bird collection from South Kensington to the site of the Rothschild Museum at Tring in Hertfordshire. Pat Hall was opposed to the move on scientific grounds. It is easy to see why. She would also have had to move from Surrey to somewhere nearer Tring. She therefore decided to give up her work at the Museum in 1971. At the same time she moved from Epsom to the New Forest in Hampshire where it was noted she devoted her time ‘to friends, dogs, horses and village life’. However, retirement from the ornithological world in which she was well known came more gradually. She was in demand as an editor, speaker and organiser. What appears to have been her last job was presidency of the 4th Pan-African Ornithological Congress in 1976. That caused her particular hassles because of its move at the last minute from Kenya to the Seychelles over political problems.

It was said that the award which gave her particular pleasure was the Stamford Raffles Award of the Zoological Society of London for 1971—presented by the Society’s President, Prince Philip, Duke of Edinburgh at the Annual General Meeting I attended in 1972. This is how the Annual Report recorded it:

The Stamford Raffles Award (awarded to an amateur zoologist for distinguished contributions to zoology) to Mrs B. P. Hall, British Museum (Natural History), in recognition of her work on the taxonomy and zoogeography of birds, particularly those of Africa. The award was a sculpture in bronze, Wild Boar, by Miss Elisabeth Frink.

It was though the final paragraph of citation for the Union Medal of the British Ornithologists’ Union that best sums up what she had achieved:

Her scientific work has already been widely recognised, by her election as Corresponding Fellow of the American Ornithologists' Union, and by the awards of the Gill Memorial Medal of the South African Ornithological Society and the Stamford Raffles Award of the Zoological Society of London. It is most fitting that the Union Medal should now be awarded to Pat Hall, who has so strikingly shown that the heights  of ornithological achievement can still be scaled without the benefit of formal academic qualifications and institutional backing.

Retirement saw her adding two books to the one she had written with Derek Goodwin in 1969, a book published privately of nonsense verse about their lives in the Museum, Bird Room Ballads; Alfred Hughes provided the illustrations. The next was the story of her life in the Mechanised Transport Corps, What a Way to Win a War (1978, Midas Books, Tunbridge Wells). Finally, in 1993 came A Hawk from a Handsaw (1993, privately published). I have not been able to find a copy of the latter but her obituarist for The Ibis noted that it gave her side of what proved to be an unhappy collecting trip to Angola.

I wonder if Alfred Hughes realised that in speaking to his friend Kinnear he was launching his cousin Pat Hall on a pathway to her becoming a major player in classical ornithology of the 20th century. The Hughes-Hall family axis runs wide and deep in the ornithology of Africa and Asia.

Beryl Patricia Hall died on 4 August 2010 in a nursing home at Fishbourne, Hampshire.

Prys-Jones R. 2011. Beryl Patricia Hall (née Woodhouse), 1917-2010. Ibis 153, 913-914.

UPDATED 24 January 2025

An American Crocodile: Honduras 2024

Last month we saw this American Crocodile (Crocodylus acutus) in the lower reaches of the Rio Cuero, Honduras, just about in the centre of its geographical range.

The American Crocodile prefers the somewhat saline waters of estuaries and mangroves and in this respect resembles the Estuarine or Saltwater Crocodile (C. porosus) of south-east Asia and northern Australia, a story I will return to in my Salt Glands Revisited series. It is also one of the largest species of crocodilian.

Basking with the mouth open—and thereby showing their very impressive set of teeth—appears to be concerned mainly with thermoregulation. Under the sun the head warms more quickly than the bulky body. Evaporative cooling from the tissues lining the mouth will lower the temperature of venous blood leaving the head. Provided there is a countercurrent arrangement whereby arterial blood entering the head is cooled by the venous blood passing in the opposite direction the brain will be kept relatively cool while the rest of the body gets to the optimum temperature. That’s the standard story but I have not checked whether or not there has been any experimental demonstration of the cooling in action. The reason I wonder, is that if, during basking, the blood from the body is relatively cool why the high rate of blood flow to the brain does not also keep the brain cool without the need for evaporative cooling? Or does the head get so hot that cooler blood from the body is insufficient to prevent the brain from suffering heat damage? Somebody who knows a lot more about crocodiles and their thermal physiology will put me right.

We asked if the crocodiles were dangerous to the human population especially the children. We were told not. Salties or Nile Crocodiles they are not.

George Cuvier in 1807 was the first to describe the American Crocodile but considered it another species of alligator. It took another Frenchman, the brilliant but mercurial Constantine Samuel Rafinesque, then working in the USA, to suggest in 1822 that it was not alligator but a crocodile.

Crocodylus acutus Distribution
Achim Raschka
BB BY-SA 3.0 Wikimedia Commons

Dear Sam, Sorry to hear you died in 1956. The crassness of modern ‘tech’

The crassness of modern life is sometimes exemplified by the ‘tech’ companies that make their business the distribution of scientific papers from the past and present and an attempt to be the site of discussion between scientists.

I was sent a link by one of their regular and too frequent emails drawing my attention to a paper by Samuel Brody (with Arthur Chester Ragsdale, 1890-1969, and Charles Wesley Turner, 1897-1975). The last time I actually read that paper was as an old-fashioned pre-Xerox photocopy made in the 1950s in the collection of a late colleague. How nice to read it again and within milliseconds it was downloaded. But then came a the follow-up seconds later inviting me to thank the author and to tell him what had sparked my interest in the paper. There was though a problem. The author had died in 1956 so where was the message to be sent? The same place as emails to Santa Claus?

It is difficult to appreciate the mindset of those who think the science they are interested in began within the last ten years. Especially that is when they are encouraged to think so by PhD supervisors, as, regrettably, is the case even in UK here we once had a more rigorous respect for knowing the literature.

So I shall not be writing to thank Sam Brody but it is worth pointing out who he was and why the research he did is still important.

Sam Brody ca 1930 when he was a
Guggenheim Fellow in 1929 and 1931
From here

Sam Brody produced the famous mouse-to-elephant curve which shows how metabolic rate varies with body weight and which followed up Max Kleiber’s work on a few species which suggested that metabolic rate (effectively oxygen consumption) did not increase linearly with body eight but as around the three-quarter power of body weight. In other words metabolic rate per kilogram is much lower in the elephant than it is in the mouse. The reason for this relation between metabolic rate and body weight is still a very active interest although it does seem that a present more heat has been generated than light has been shed on the problem in recent years.

A second area of research in which Sam Brody is remembered is for his demonstration that the frequency of milking in dairy animals leads to an increase in the rate of milks secretion. The mechanism by which that happens kept me occupied for many years and has particular application in not only dairy animals but also in human lactation where the frequency and completeness of milk removal are key determinants of breastfeeding success.

Samuel Brody was professor of dairy husbandry in the University of Missouri. He was born in Lithuania in 1890. He emigrated to Canada in 1906 where he worked as a machinist, miner, fisherman and door-to-door salesman. He then obtained a place across the border as a student at the National Agricultural School in Pennsylvania. Hearing of great things happening at the University of California at Berkeley he had to teach himself to read and write in English in order to pass the entrance exam and study there. He graduated in 1917 in biochemistry. Following a masters he spent a short time in the US Air Force and worked in the University of California until he appointed to his post in Missouri in 1920. He died in his office in August 1956.

Brody’s work on growth, metabolism, thyroid function and milk production established his international reputation—which leads me on to my second point. It is little recognised that advances in human reproductive physiology in particular and in human health generally have been made under the aegis of funding for agricultural research. With that funding drastically reduced in recent decades, particularly in UK, that generation of new knowledge has ceased—yet another reason for condemning successive governments and devolved administrations for following the mantra of its unimportance with agriculture only being responsible for 2% of GDP but ignoring the rather important matter of national food security and its 100% role in keeping people alive.

Meanwhile the crassness continues. In requesting that I ask co-authors to join, a list is provided of those not belonging. That list includes one who has been dead for 49 years.

Long-tailed Skink in Hong Kong

 

This superb Long-tailed Skink (Eutropis longhicaudata) joined AJP for luch in Victoria Park, Hong Kong, last week. It is the largest skink that occurs in Hong Kong, reaching 40 cm in length. It occurs in south-east Asia, from Malaysia in the south to southern China and Taiwan in the north.

Salt Glands Revisited. 3. Maryanne Robinson Hughes (1930-2020)

Fifty years ago the late Jim Linzell and I were writing our monograph, Salt Glands in Birds and Reptiles, for the Physiological Society's Monograph Series; it was published in May 1975. In this series I revisit some of the topics and people who followed up the discovery of salt glands in birds by Knut Schmidt-Nielsen.

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I was sad to find that Maryanne Robinson Hughes had died in 2020 just short of her 90th birthday. Maryanne was in right at the beginning of the discovery of salt glands in birds since she was Knut Schmidt-Nielsen’s PhD student at Duke University in North Carolina and in 1958 was a co-author of the second full paper, with Knut and Ragnar Fänge, to appear on salt glands in birds in which they established the nervous control of the gland.

An obituary online written by one of her three children, together with a page on ancestry.com provides an outline of her life. Maryanne Elizabeth Robinson was born on 27 December 1930 in Binghampton, New York. She was educated at Binghampton Central High School and Harpur College. A masters and PhD at Duke then followed. Her PhD was awarded in 1962 but from newspaper articles on her engagement it would seem that while writing up she had been employed as a research assistant in the physiology department of the University of Washington in Seattle. On 26 August 1961 she married Gilbert C Hughes (1933-2010) in New York; they went to live in Kansas where Gil was an assistant professor. In 1964 the moved across the Canadian border, Gil to the Department of Botany of the University of British Columbia in Vancouver where he worked on marine fungi. I remember Maryanne telling me that with a young family she first worked part-time in the Department of Zoology where she stayed until retirement. Maryanne died in a care home in Vancouver on 10 December 2020.

I met Maryanne only once; at a dinner with Gil, during the Physiological Congress on the UBC Campus in Vancouver, where I chaired a session on salt glands, a memorable occasion (the dinner not the session) with them both in fine form. Maryanne continued working on birds with salt glands for her entire career with a string of postgraduate students and other collaborators. She was particularly concerned with the integration of function of all the organs and tissues concerned with osmoregulation and water balance, a very difficult job given the nature of urinary excretion and the admixture of urine and faeces in the hindgut of birds.

I have gathered Maryanne’s publications in the following list from my own records and all the usual sources. Her first paper was published when she was 27; her last when she was 76. However, because the indexing services often miss (even with the benefit of machine-learning) many publications in books and conference proceedings, the list may not be complete. Some of Maryanne’s papers in my own records have not been picked up.

Maryanne Robinson Hughes: Publications 1958-2007 

Fänge R, Schmidt-Nielsen K, Robinson M. 1958. Control of secretion from the avian salt gland. American Journal of Physiology 195, 321-326.

Hughes MR. 1962. Studies on renal and extrarenal salt excretion in gulls and terns. PhD thesis, Duke University, North Carolina.

Goldstein DL, Hughes MR, Braun EJ. 1966. Role of the lower intestine in the adaptation of gulls (Larus glaucescens) to sea water. Journal of Experimental Biology 123, 345-357.

Hughes MR. 1968. Renal and extrarenal excretion in the common tern Sterna hirundo. Physiological Zoölogy 41, 210-219.

Hughes MR, Ruch FE. 1968. Sodium and potassium in the tears and salt gland secretion of saline acclimatized ducks. Proceedings of the International Union of Physiological Sciences 7, 204 (24th International Congress, Washington, D.C.).

Hughes MR. 1969. Ionic and osmotic concentration of tears of the gull, Larus glaucescens. Canadian Journal of Zoology 47, 1337-1339.

Hughes MR, Ruch FE. 1969. Sodium and potassium in spontaneously produced salt-gland secretion and tears of ducks, Anas platyrhynchos, acclimated to fresh and saline waters. Canadian Journal of Zoology 47, 1133-1138.

Hughes MR. 1970. Flow rate and cation concentration in salt gland secretions of the glaucous-winged gull, Larus glaucescens. Comparative Biochemistry and Physiology 32, 807-812.

Hughes MR. 1970. Relative kidney size in nonpasserine birds with functional salt glands. Condor 72, 164-168.

Hughes MR. 1970. Some observations on ion and water balance in the puffin, Fratercula arctica. Canadian Journal of Zoology 48, 479-482. 

Hughes MR. 1970. Cloacal and salt-gland ion excretion in the seagull, Larus glaucescens, acclimated to increasing concentrations of sea water. Comparative Biochemistry and Physiology 32, 315-25. 

Hughes MR. 1972. The effect of salt gland removal on cloacal ion and water excretion in the growing kittiwake, Rissa tridactyla. Canadian Journal of Zoology 50, 603-610. 

Hughes MR. 1972. Hypertonic salt gland secretion in the glaucous-winged gull, Larus glaucescens, in response to stomach loading with dilute sodium chloride. Comparative Biochemistry and Physiology A 41. 121-127.

Hughes MR, Blackman JG. 1973. Cation content of salt gland secretion and tears in the Brolga, Grus rubicundus (Perry) (Aves: Gruidae). Australian Journal of Zoology 21, 515-518.

Hughes MR. 1974. Water content of the salt glands and other avian tissues. Comparative Biochemistry and Physiology A 47, 1089-1093. 


Hughes MR. 1975. Salt gland secretion produced by the gull, Larus glaucescens in response to stomach loads of different sodium and potassium concentration. Comparative Biochemistry and Physiology A 51, 909-913.

Ruch FE Jr, Hughes MR. 1975. The effects of hypertonic sodium chloride injection on body water distribution in ducks (An as platyrhynchos), gulls (Larus glaucesens) and roosters (Gallus domesticus). Comparative Biochemistry and  Physiology A 52, 21-28.


Hughes MR. 1976. Effect of glucose on salt gland secretion in the glaucous-winged gull, Larus glaucescens. Comparative Biochemistry and Physiology A 53, 311-312. 

Hughes MR. 1976. The effects of salt water adaptation on the Australian black swan, Cygnus atratus (Latham). Comparative Biochemistry and Physiology A 55, 271-277.

Hughes MR. 1977. Observations on osmoregulation in glaucous-winged gulls, Larus glaucescens, following removal of the supraorbital salt glands. Comparative Biochemistry and Physiology 37A, 281-287.

Walter A, Hughes MR. 1978. Total body water volume and turnover rate in fresh water and sea water adapted glaucous-winged gulls, Larus glaucescens. Comparative Biochemistry and Physiology 61A, 233-237.

Hughes MR. 1980. Glomerular filtration rate in saline acclimated ducks, gulls and geese. Comparative Biochemistry and Physiology 65A, 211-213.

Roberts JR, Hughes MR. 1983. Glomerular filtration rate and drinking rate in Japanese Quail, Coturnix coturnix japonica, in response to acclimation to saline water. Canadian Journal of Zoology 61, 2394-2398.

Roberts JR, Hughes MR. 1983. The effect of saline acclimation on water and sodium transport across the small intestine of ducklings, gulls and quail. American Zoologist 23, 1014.

Roberts JR, Hughes MR. 1984. Exchangeable sodium pool size and sodium turnover in freshwater- and saltwater-acclimated ducks and gulls. Canadian Journal of Zoology 62, 2142-2145.

Roberts JR, Hughes MR.  1984. Saline acclimation and water and sodium transport across avian small intestine. American Journal of Physiology 247, R246-249. 

Hughes MR. 1987. The effects of ureteral resistance on gull urine composition and flow rate. Canadian Journal of Zoology 65, 2669-2671.

Hughes MR, Roberts JR, Thomas BR. 1987. Total body water and Its turnover in free-living nestling glaucous-winged gulls with a comparison of body water and water flux in avian species with and without salt glands. Physiological Zoology 60, 481-491.

Conway GL, Hughes MR, Moldenhauer RR. 1988. Extrarenal salt excretion in clapper and king rails. Comparative Biochemistry and Physiology A 91, 671-674.

Hammons RL, Hughes MR, Moldenhauer RR. 1988. Body water and water flux in fresh water and sea-water acclimated clapper rails, Rallus longirostris. Comparative Biochemistry and Physiology A 91, 539-541.

Hughes MR, Roberts JR, Thomas BR. 1989. Renal function in freshwater and chronically saline-stressed male and female Pekin ducks. Poultry Science 68, 408-416.

Hughes MR. 1989. Extracellular fluid volume and the initiation of salt gland secretion in ducks and gulls. Canadian Journal of Zoology 64, 194-197.

Hughes MR, Chadwick A. (Editors) 1989. Progress in Avian Osmoregulation. Leeds: Leeds Philosophical and Literary Society.

Hughes MR. 1989. Stimulus for avian salt gland secretion. In, Progress in Avian Osmoregulation. Edited by MR Hughes and A Chadwick, pp 143-161. Leeds: Leeds Philosophical and Literary Society.

Hughes MR, Kasserra C, Thomas BR. 1990. Effect of externally applied bunker fuel on body mass and temperature, plasma concentration, and water flux of Glaucous-winged Gulls, Larus glaucescens. Canadian Journal of Zoology 68, 716-721.

Hughes MR, Winkler D. 1990. Osmoregulation in nestling California gulls at Mono Lake, California. Comparative Biochemistry and Physiology A 95, 567-571.

Kasserra CE, Jones DR, Hughes MR. 1991. Acid-base disturbance and ventilatory response to changes in plasma osmolality in Pekin ducks. Respiration Physiology 85, 383-393.

Hughes MT, Zenteno-Savin T, Kojwang D. 1991. Effects of saline acclimation and cecal ligation on body water and water flux in male and female Pekin ducks. Canadian Journal of Zoology 69, 771-775.

Hughes MR, Kojwang D, Zenteno-Savin T. 1992. Effects of caecal ligation and saline acclimation on plasma concentration and organ mass in male and female Pekin ducks, Anas platyrhynchos. Journal of Comparative Physiology B 162, 625-631.

Bennett DC, Bowes VA, Hughes MR, Hart LE. 1992. Suspected sodium toxicity in hand-reared great blue heron (Ardea herodias) chicks. Avian Diseases. 36, 743-748. 

Kojwang D, Hughes MR. 1993 High dietary sodium chloride and body temperature in the domestic fowl and the glaucous-winged gull. Journal of Comparative Physiology B 163, 421-426.

Hughes MR, Goldstein DL, Raveendran L. 1993. Osmoregulatory responses of glucous-winged gulls (Larus glaucescens) to dehydration and hemorrhage. Journal of Comparative Physiology B. 163, 524-31. 

Hughes MR. 1995. Responses of gull kidneys and salt glands to NaCl loading. Canadian Journal of Physiology and Pharmacologu 73, 1727-1732.

Bennett DC, Hughes MR, De Sobrino CN, Gray DA. 1997. Interaction of osmotic and volemic components in initiating salt-gland secretion in Pekin ducks. Auk 114, 242-248.

Ching AC, Hughes MR, Poon AM, Pang SF. 1999. Melatonin receptors and melatonin inhibition of duck salt gland secretion. General and Comparative Endocrinology 116, 229-240.

Bennett DC, Hughes MR, Elliott JE, Scheuhammer AM, Smits JE. 2000. Effect of cadmium on Pekin duck total body water, water flux, renal filtration, and salt gland function. Journal of Toxicology and Environmental Health A. 59, 43-56.


Hughes MR, Smits JE, Elliott JE, Bennett DC. 2000. Morphological and pathological effects of cadmium ingestion on Pekin ducks exposed to saline. Journal of Toxicology and Environmental Health A. 61, 591-608. 

Hughes MR, Bennett DC, Gray DA, Sharp PJ, Scheuhammer AM, Elliott JE. 2003. Effects of cadmium ingestion on plasma and osmoregulatory hormone concentrations in male and female Pekin ducks. Journal of Toxicology and Environmental Health A 66, 565-579. 

Hughes MR. 2003.Regulation of salt gland, gut and kidney interactions. Comparative Biochemistry and  Physiology A 136, 507-524. 

Bennett DC, Hughes MR. 2003. Comparison of renal and salt gland function in three species of wild ducks. Journal of Experimental Biology 206, 3273-3284. 

Bennett DC, Gray DA, Hughes MR. 2003. Effect of saline intake on water flux and osmotic homeostasis in Pekin ducks (Anas platyrhynchos). Journal of Comparative Physiology B. 173, 27-36.

Bennett DC, Kojwang D, Sullivan TM, Gray DA, Hughes MR. 2003. Effect of saline acclimation on body water and sodium compartmentalization in Pekin ducks (Anas platyrhynchos). Journal of Comparative Physiology B. 173, 21-26.

Hughes MR, Bennett DC. 2004. Effect of saline intake, sex and season on Pekin duck osmoregulatory organ masses and comparison with wild Mallards. Canadian Journal of Zoology 82, 30-40.

Bennett DC, Gray DA, Sharp PJ, Hughes MR. 2005. Redistribution of extracellular water and sodium may contribute to saline tolerance in wild ducks. Physiological and Biochemical Zoology 78, 447-455.

Hughes MR, Bennett DC, Gray DA, Sharp PJ, Poon AM. 2006. Influences of sex and saline intake on diurnal changes in plasma melatonin and osmoregulatory hormones of Pekin ducks (Anas platyrhynchos). General Comparative Endocrinology 149,124-133 

Hughes MR, Kitamura N, Bennett DC, Gray DA, Sharp PJ, Poon AM. 2007. Effect of melatonin on salt gland and kidney function of gulls, Larus glaucescens. General and Comparative Endocrinology 151, 300-307. 

William Rowan—Pioneer of Photoperiodism. 3. The horrific death of a schoolboy entomologist

Oak Eggar Moth Lasiocampa quercus
It will become obvious on reading this article why a
photograph of this species is an appropriate one to
remember the subject
From Wikipedia ©Entomart

While William Rowan was looking for a way to make a zoological living he worked a schoolmaster, latterly for a school year at Bedales in 1918-19.

Marianne Ainley in her biography wrote that Rowan arrived at Bedales School in Hampshire on 19 September 1918. She found in Rowan’s notes an account of the tragic death of a boy at the school:

…He liked all his colleagues and many of his pupils. He felt a special affinity for Woolacott, one of the big boys, a clever, reserved lad, keen on natural history, who had already published notes in the Entomologist. The two spent much time together, and from their discussions Rowan found the boy was practically a misogynist. This, he noted, "seems unnatural and calls for a deeper explanation.” Because of his busy schedule, however, Rowan had to postpone investigating the causes of the boy's attitude.
     Understandably, Rowan was distressed when, a few weeks later, Woolacott informed him he would probably be expelled. Rowan promptly arranged to see the headmaster [John Haden Badley who had, with his wife, founded the school] to intercede on Woolacott's behalf, but was too late. Woolacott shot himself that afternoon. Rowan blamed himself for not getting to the root of the boy's problem, and was most deeply touched to find that Woolacott had left a pile of beautifully made entomological slides, representing many hours of careful work, on Rowan’s desk.

Who was this budding entomologist, Woolacott? And what did reports of his death have to say?

First though it is worth pointing out that Bedales was a rare example at the time of a co-educational secondary school and I take it that the ‘misogyny’ on the part of Woolacott, was related to the presence of female pupils at the school and, perhaps, somewhat of a tradition of the boys who had objected to the presence of girls when they first appeared on the scene some years earlier.

It must also be borne in mind that the tragedy occurred on 4 November 1918, only six weeks after Rowan’s arrival. He had clearly got to know the boy’s interest in the natural world but had not had time to understand much of Woolacott’s past.

The Hampshire Telegraph of Friday 8 November had a short account of the coroner’s inquest held the day before.

STUDENT OF 17 SHOT DEAD.

     A shocking tragedy occurred at Bedales School, Petersfield, on Monday afternoon, when a student named Harold Kingsley Woolacott, aged 17, was found dead in a dormitory with a bullet wound in his temple.
     It appears that the deceased lad was about to leave the school, and on going to pack up his clothes we given a small rifle, which the Matron had been keeping for him for about two years. Shortly afterwards he was found as described with the rifle at his feet. Dr. Brownfield was summoned, but could only pronounce life extinct.
     At an inquest yesterday afternoon a verdict of "Suicide whilst temporary insane" was returned...

The school magazine, the Bedales Record for 1918-19 also had an account:

The term was saddened by the death of Woolacott, as told on a later page. He was buried in Steep Churchyard, where now lie several of our boys, the coffin being carried by his fellow prefects and followed by his schoolfellows.

H. K. Woolacott joined the School in 1916 at a later age than most. He was a quiet introspective boy, given to melancholy moods, which were deepened by the absence of his parents in India and by the war. He would normally have left the School last Summer to begin his military service, but asked to stay on another term in order to be a prefect and try to do more for the School than he had done. But he did not prove strong enough to be in so responsible a position, and at the beginning of November it was decided, with his own concurrence, that he should leave then and begin his training at once. While packing his things, amongst them a miniature rifle which, like many others, he had brought here for practice on the range, and for which, unknown to us, he had some cartridges of his own, he must, in a moment's impulse, have shot himself through the head; when found, a few moments later, he was past all help. At the inquest neither the evidence nor his own diary could throw any light upon the motive, other than the boy's strange nature and his sense of failure at School, and no kind of blame was felt to attach to anyone else.

I would suggest that Rowan’s more succinct explanation for Woolacott’s departure from the school is the correct interpretation; he was being expelled and thus required to join the army seven days, as it happened, before the Armistice of 11 January.

Harold Kingsley Woolacott had indeed had notes published in The Entomologist. I have found two, both in the January 1918 edition.

In the early decades of the 20th century amateur entomology and especially lepidoptery was in its heyday.
Lasiocampa quercus is the Oak Eggar Moth. Melitaea aurinia, the Marsh Fritillary Butterffly, is now Euphydryas aurinia. Phragmatobia fuliginosa is the Ruby Tiger moth. Macrothylacia rubi is the Fox Moth. Manulea lurideola is the Common Footman Moth

As mentioned by the report in the school magazine Harold Woolacott’s parents were in India. He was born in Brixton, London in 1901 to John Evans Woolacott and his second wife, Angiolina Maria Emily Seneca. John Evans Woolacott (1861-1936) is described on his Wikpedia page as journalist, newspaper editor and political activist who worked first in London for the Central News Agency as lobby correspondent and then in Ireland, Morocco and Egypt. in 1895 he stood, unsuccessfully, for parliament as a member of the Independent Labour Party in the Rollox Division of Glasgow. During the 1890s he worked on The Democrat and was assistant editor of the Weekly Dispatch. His first wife died and he married Angiolina Seneca in 1897; he was 34 and she was 21. By 1903 he was assistant editor of The Economist. In 1908 he was elected president of the Institute of Journalists but then left for India where he was assistant editor of The Statesman in Calcutta. 1913 saw him as editor of the Bombay Gazette. That publication soon closed and he returned to UK. He swapped political allegiance being adopted as a prospective parliamentary candidate for the Liberal Party but because of the outbreak of war no general election was held. Woolacott returned to India in 1916 (the year his son was sent to Bedales) working for The Pioneer in Allahabad and as a correspondent for The Times of London. By the time he had returned to UK in 1925 his politics had changed again, writing books opposed to political change in India. In 1929 he was appointed editor of The Bioscope, a weekly magazine devoted to the cinema. John Evans Wollacott died in Surrey in 1936.

Harold Woolacott’s mother, Angiolina, was the daughter of Alfonso and Eliza Seneca. Alfonso Seneca was Italian, a professional singer who appears in the local London newspapers of 1877 and 1878 performing regularly and frequently in concerts. Alfonso died in 1880. As far as I can see from the records John and Angiolina had no other children. Angiolina died in Surrey in 1934.

I have been able to find nothing else about the unhappy and clearly disturbed Harold Kinglsey Woolacott. I do not know if he ever accompanied his parents to India. It is clear that he entered Bedales when his parents left UK for the second time but where had he been educated until then? It was the norm for children of parents in India to either send or leave their children in UK when they were seven. The only clue as to young Woolacott’s location at any time is the address given in one of the articles in The Entomologist; in the summer of 1917 he was in St Merryn in Cornwall.

I have found a family tree online which includes a brother and sisters of John Evans Woolacott but not including John Evans himself. I do not know if Harold would have been in touch with members of his extended family while his parents were in India.

Harold Kingsley Woolacott’s tombstone can be found in the graveyard of All Saints in the village of  Steep, near Petersfield. Reading his notes to The Entomologist I suspect the budding entomologist was a young scientist in the making. I can see why William Rowan, fondly remembered for his teaching and encouragement of birdwatching and photography in the school, was so upset by Woolacott’s utterly tragic death 106 years ago today.

Gravestone of
Harold Kingsley Woolacott
from findagrave.com

Ainley MG. 1993. Restless Energy. A Biography of William Rowan 1891-1957. Montreal: Véhicule Press.

Pheasant-tailed Jacana in Hong Kong

 

AJP spotted this Pheasant-tailed Jacana (Hydrophasianus chirurgas) at Nam Sang Wai two weeks ago, his third ever. It is classed as an uncommon migrant and rare winter visitor.  This is in non-breeding plumage, a time of year when the long pheasant-type tail is absent. The only migratory jacana they occur from Yemen in the west to the Philippines in the east.

British Journal of Herpetology (1948-1985). All issues available online with free access

I was delighted to see that the British Herpetological Society now has all issues of British Journal of Herpetology available with free access on its website. This is exactly the way a proper learned society should act and is in marked contrast to commercial scientific journal publishers and some scientific societies who know no better than to exist as parasites on the acquisition and dissemination of knowledge.

British Journal of Herpetology was launched by the newly formed society in 1948 and ran under that title until shortly after I handed over the editorship of the journal to Trevor Beebee in 1985. The change of title—to Herpetological Journal—had as I recall two aims: to ensure that the coverage was not just perceived as British amphibians and reptiles; submissions were not restricted to authors in Britain.

Access to the papers published in BJH had never been easy because the Society had relatively few sales to libraries. The journal was distributed to members but from the early years I suspect few copies have survived.

Apart from those wishing to consult the papers themselves, the additional material, the early membership list, for example, provides a fascinating insight into the development of interest in reptiles and amphibians in Britain and into what a good job those who founded the Society did in attracting members from such a wide range of backgrounds and interests—a topic I will return to in future articles.

A Snipe but not a Common Snipe in Hong Kong

 

AJP spotted this snipe at Nam Sang Wai last week. It is either, as confirmed by that very co-operative underwing view, a Pin-tailed Sbnipe (Gallinago stenura) or a Swinhoe’s Snipe (G. megala). Unfortunately, only with a bird in the hand and the tail feathers spread is it possible to tell t’other from which. Both are passage migrants in Hong Kong. The Pin-tailed breeds in Russia and Mongolia and winters from India to northern Australia. Swinhoe’s breeds in central Asia, Mongolia and Russia and winters across southern Asia to Australasia.

Salt Glands Revisited. 2. We should have mentioned the other glands that drain into the nose

Fifty years ago the late Jim Linzell and I were writing our monograph, Salt Glands in Birds and Reptiles, for the Physiological Society's Monograph Series; it was published in May 1975. In this series I revisit some of the topics and people who followed up the discovery of salt glands in birds by Knut Schmidt-Nielsen.

- - - - - - - - - - - - - - - - - - - - - - - 

One problem with writing a book 50 years ago was that the copy that arrives in the reader’s hands has a sense of finality. Last thoughts, changes of mind or new evidence can only be passed on by producing a new edition some tears later or covering the same ground in an invited review. That problem, although still present, is lessened by having a website covering a particular publication that can be amended accordingly.

One point we forgot to mention when we wrote Salt Glands in Birds and Reptiles was the possibility that secretion from other glands could perhaps be mistaken for secretion from the nasal salt glands of birds. Small drops or dampness seen within the nostril in response to cholinergic drugs or during periods of excitement could be interpreted as a sign of the salt gland being activated at a low rate or for a brief period. However there remains the possibility, particularly if the secretion that appeared was not analysed, that the small amounts could arise from other glands in the head, the secretions from which can find their way to the nostrils.

The Harderian gland is a relatively large gland that sits behind the eyeball. It is particularly associated with tetrapods which have a nictitating membrane. A number of functions of its secretion have been described or postulated including, of course, lubrication and protection, including immunoprotection, of the eye. In short, it is a second source of tears, alongside the lachrymal gland which in birds is very small. Just as when we cry, tears appear not only from the eye itself but also pass down the nasolacrimal duct to the nose; sniffling and lachrymosity go together.

We reproduced this drawing in Salt Glands in Birds and Reptiles
I have added colouring to show the nasal salt gland and the Harderian Gland

There is no indication that in birds with a nasal salt gland that the Harderian gland plays any part in removing excess salt. In geese, for example, we found no evidence for any increase in blood flow through the Harderian gland. By contrast, salt gland blood flow increased increased up to 26-fold when secretion was activated by a salt load. However, there is one intriguing report in the literature that the Harderian gland might be doing something in relation to salt and water balance in some birds.

In 1968 the late Maryanne Robinson Hughes and Frank E Ruch of the University of British Columbia noticed that in domestic ducks kept on sea water the feathers between the beak and the eye were wet and contained depressions which were filled with a clear fluid. Analysis of the fluid showed the sodium concentration to be similar to that of blood plasma. However the potassium concentration was much higher than that in plasma. Moreover, the potassium concentration increased with higher salinities up to full strength sea water, the main rising occurring when the salinity of the water exceeded that of plasma. No such effect was seen in gulls and the suggestion was that ducks living on sea water are at a precarious state of salt and water balance compared with gulls which can cope with the salinity of sea water much more readily. Although the source of the potassium-rich tears was not identified, the Harderian gland is the most likely since since the lachrymal glands are tiny, only one hundred of the weight of the Harderian glands.

Pertinent to the point of this article is that it was impossible to determine just how much potassium the Harderian glands were secreting in the ducks since tears could be passing down the nasolacrimal duct. However, they did find a band of smooth muscle around the opening of this duct which they suggested could act as a sphincter regulating the passage of tears from the eye by that route.

If only we had mentioned it at the time I would not have needed to write this article 50 years later.

Fänge R, Schmidt-Nielsen K, Robinson M. 1958. Control of secretion from the avian salt gland. American Journal of Physiology 195, 321-326.

Hughes MR, Ruch FE. 1969. Sodium and potassium in spontaneously produced salt-gland secretion and tears of ducks, Anas platyrhynchos, acclimated to fresh and saline waters. Canadian Journal of Zoology 47, 1133-1138.

Peaker M, Linzell JL. 1975. Salt Glands in Birds and Reptiles. Cambridge: Cambridge University Press.

William Rowan—Pioneer of Photoperiodism. 2. The Biography

I have enjoyed reading the late Marianne Gosztonyi Ainley’s (1937-2008) account of William Rowan’s life, from its beginning, in 1891 at Basel in Switzerland, as the son of a railway engineer from Northern Ireland and a Danish mother, to the end, in 1957, at home alone in Edmonton, Alberta.

Marianne Ainley covered all aspects of Rowan’s life in detail. Since there is available online what is essentially a précis of her 378 page book I will not repeat the basic information other than to say that Rowan had become obsessed with the idea of getting to western Canada while at Bedford School. He was inspired to travel, study, draw and photograph the wildlife, by visits to the school by Richard Kearton, showing his brother Cherry’s photographs of birds, and by Ernest Thompson Seton who talked about the large mammals of Canada and the USA. Rowan’s widowed mother held and the pursestrings and was eventually persuaded and she arranged for him to be a ranch pupil with a British family in Alberta. Thus Rowan was in Alberta for a first time in 1908, to the less than comfortable world of a ranch pupil but travelling first class on his mother’s insistence. He returned to Britain in 1910 where he crammed for matriculation to London university. He then returned to Canada, doing the odd job in Winnipeg while roaming and photographing the country and its wildlife. He was persuaded both by friends in Canada and family to take the place at University College London; he then had to find his own fare for the voyage.

The Dark-eyed Junco (Junco hyemalis) was one of two
species used in early experiments on varying daylength*

Thus began Rowan’s the zoological career but in describing it Ainley left a number of unanswered questions while missing clues that could have led to explanations of events. Some of Ainley’s interpretations are amusingly wrong while others are completely out of touch with life, the university scene or science in the early years of the 20th century. On a general note I often find accounts written by historians of science to be unsatisfactory simple because those historians have not been steeped in the discipline or of how practitioners within that discipline lived and operated. Similarly, conventions of the time are often judged through the lens of the present. Thus it was not at all strange that Mrs Rowan was left at home to do the housework and look after the children. Nor was it, or is it, odd that Rowan in following his many interests neglected to give his wife and children his full attention. The response of most of my scientific acquaintances to that statement would be, ‘Yes…and?’

Rowan clearly enjoyed studying zoology and botany, particularly the field work involved, while failing intermediate physics twice. He also wrote popular articles and took photographs while a student. With the course uncompleted he, like many of his fellow students, enlisted in the 14th (Reserve)  County of London Battalion of  London Scottish Regiment, a month after the declaration of war with Germany. However, Private Rowan became ill during his first long leave, during which time he returned to the labs at UCL. He had bouts of illness throughout his life. His leave was extended and the army then discharged him as ‘no longer fit physically for war service’. He had served 1 year, 68 days when he returned to civilian life in November 1915. With the family fortunes, which were held mainly in French stocks and shares, declining because of the war, Rowan spent a year in pursuing energetically his interests in natural history, while worrying about getting a job such while considering lecturing and taxidermy as possibilities. In October 1916 he returned to UCL for his much delayed final year. He graduated in 1917 with Third Class Honours, six years after he had begun life as a student.

Although not commenting specifically on the class of his honours degree, Ainley argued that Rowan never really got the hang of doing exams and that Bedford School was more interested in producing sportsmen than learned gentlemen. While later in the century, a ‘third’ would have been a bar to further life in a university, this was clearly not the case at the time.

Rowan spent a over year teaching at Eastby and at Bedales, a public (i.e. for non-British readers, a non-state, fee-paying, private school) school in Hampshire. Then came the offer of several jobs and his acceptance of one of them, assistant in the Department of Zoology at UCL, in effect an assistant lectureship including responsibilities within the departmental museum. He had, however, through a contact, applied for a new post in the University of Manitoba in Winnipeg. Keen to return again to Canada, its wild places and its wildlife, he amassed a great deal of support for his application from leading British zoologists:

[J.P>] Hill stressed Rowan's qualifications in zoology, his wide knowledge of ornithology, his artistic talent, his excellence as a lecturer, and his "driving power and considerable organising ability.” Oliver wrote about Rowan's resourcefulness as a naturalist and his knowledge of bird protection. Dr. P. Chalmers Mitchell, secretary of the Zoological Society of London, praised Rowan's aptitude for fieldwork and his capacity for executive work.

He was offered the post in Winnipeg and therefore had to resign from the UCL job before he had started.

A month before he left for Canada in October 1919, Rowan married Reta Guenever Mary Bush whom he had met while she was an art student at the Slade. From their arrival in Quebec, they made their way to Winnipeg. After a short time in Winnipeg, Rowan was recruited to the University of Alberta by Tory. The pay was better with a promise of a chair in the offing.

Rowan’s work on the importance of changing daylength as the trigger for migration was done on a shoestring and his ‘Restless Energy’, the title of Ainley’s biography, meant that had had lots of other activities in hand, often out in the wilds of western Canada, in addition to starting a department from scratch including gathering and preparing specimens and posters. He worked at home, in the lab (such as it was) and in the field for virtually every hour of every day. However, it is obvious that he seemed unable to prioritise, leaving his daylength experiment to a helper over a crucial period, for example.

His friends in UK to whom he complained about Tory’s animus, lack of funding and lack of appreciation of his work in Canada really could not understand why Rowan stayed in Alberta. Indeed after his death Julian Huxley stated that ‘Rowan was one of the best experimental zoologists of the 20th century. But why, with all his talents, did the fool have to bury himself in Alberta?”

The American Crow (Corvus brachyrhynchos) played
a major part in migration experiments in the wild** 

Given his association with Huxley it seems anomalous that he, at one stage in his life, was supporting Lamarckism and hoped that with his crow experiments in which birds were exposed to different daylengths ‘he may be able to adduce some evidence in support of the Lamarckian concept of evolution’.  Although, as Ainley remarked, belief in Lamarck was not unusual amongst confused zoologists of the 1920s, the year the newspapers reported this, 1931, is interesting and Ainley seems to have missed the significance. In 1931 Rowan was persuaded by his friends to apply for the chair of zoology at McGill. He assembled a body of supporters in Britain and the USA. Amongst them was Ernest MacBride, a previous professor of zoology in McGill and now at Imperial College in London. MacBride, as I have recounted here was both an ardent Lamarckian, denying the existence of genes or mutations, and extreme eugenicist. MacBride has been described as Lamarck’s last disciple on earth and it cannot have done Rowan any harm in MacBride’s eyes to be seen as a Lamarckist. However, whether or not Julian Huxley, one of those responsible for the great synthesis of genetics, development and morphology in Darwinian evolution, got to know of Rowan’s statements I do not know. Despite the enormous level of support, Tory managed to kill off any chance of Rowan getting the McGill chair. In Alberta he stayed.

With the level of support given by leading zoologists in Britain and his international reputation, I find it odd that Ainley made no mention of a missing accolade. William Rowan was not a Fellow of the Royal Society. Had he been proposed but not elected? Or had his putative proposers not been able to garner enough support for a proposal to be made?

One might have thought that once the active, local, torment of Tory had disappeared off the scene and that once he finally made it to full professor (1931) he would have had a relatively untroubled existence in Alberta, even if money remained in short supply for his research. However, as he got closer to retirement long-standings members of his own department—whom he had backed relentlessly for advancement and better pay—turned against him; a demonstration of ‘no good turn goes unpunished’.

Given the explosion of interest in the photoperiodic control of reproduction that followed in Rowan’s wake it is noticeable how poorly or insufficiently appreciated in their own countries were some of its most successful scientists, ‘Jock ‘Marshall is one example; Don Farner (1915-1988) another.

By the time of Rowans death in 1957, the world of photoperiodism had moved on to the role of day length in controlling the onset of reproduction, rather than migration, and to the burgeoning field of neuroendocrinology.  Rowan’s work was known about but no longer referred to. For example, in three long reviews of aspects of environmental control of reproduction in birds and the physiological mechanisms responsible for a symposium I organised in 1973, Rowan did not get a mention.

Despite my qualms, some of which I have pointed out, Marianne Ainley made a great contribution to our knowledge of William Rowan. She, herself, had a remarkable life history. Born Marika Veronika Gosztonyi in Budapest in 1937, she  escaped from the Russian tanks advancing to put down the revolution of 1956 by walking along the railway line into Austria. From there she was able to reach an uncle in Sweden. In 1958 she moved to Canada and worked as a technician and research assistant having studied chemistry in Budapest for four years. A keen birdwatcher, she changed tack completely and became a research assistant in the history of science. In that field she obtained an MSc and then a PhD at McGill on the history of ornithology and avian biology in Canada. She was then at Concordia University and finally, as a full professor at the University of Northern British Columbia. She retired in 2002 and died of cancer in 2008.

I leave William Rowan in the words with which Marianne Ainley introduced Restless Energy:

WILLIAM ROWAN has been considered a Renaissance man, a man of integrity, a famous biologist, a flamboyant showman, a challenging teacher, and often a nuisance. His work as a scientist has been highly regarded by biologists all over the world, and his experiments have been covered in numerous zoology textbooks. Who was this scientist? What was he like as a person, a family man, a colleague, a friend? Why was he so well known to contemporary scientists, and why do we still find written references to his scientific work while, apart from anecdotal descriptions about his escapades in the field, little is known about the details of his life?

Convinced that William Rowan was an important subject for a biography, I went to Edmonton in July 1985 to see the environment where he lived and worked; to talk to the people who knew him as friend, teacher, and colleague, and who admired him as a world-renowned scientist, highly respected wildlife artist, and outstanding conservationist. Inevitably, other facets of his character came to light: his integrity, his kindness, his egotism, his unfailing need to criticize mediocrity, his ability to inspire others, his love of beauty and of nature. I learned of his exquisitely designed conservation stamps, Christmas cards, and hand-painted menus; of his great skill as a self-taught sculptor and musician, and as a builder of model sailing boats and a unique single-gauge model railway. I found that he had been well known as a radio personality, and as the man who tried to make crows fly the wrong way. Nearly thirty years after his death, William Rowan was still very much alive in Edmonton, and even those who had never set eyes on him could tell a few choice stories about his escapades. Indeed, as I gradually discovered, Rowan was an unusual individual — a Renaissance man in Alberta.

*https://commons.wikimedia.org/wiki/File:Dark-eyed_Junco,_Washington_State_02.jpg

** CC BY-SA 3.0, https://commons.wikimedia.org/w/index.php?curid=2310490

Ainley MG. 1988. Rowan vs Tory: Conflicting views of scientific research in Canada, 1920-1935. Scientia Canadensis: Canadian Journal of the History of Science, Technology and Medicine 1, 3-21.

Ainley MG. 1993. Restless Energy. A Biography of William Rowan 1891-1957. Montreal: Véhicule Press.

Houston CS. 2009. In Memoriam: Marianne G Ainley, 1937-2008. The Auk 126, 699.

 

William Rowan—Pioneer of Photoperiodism. 1. His persecutor

 It must have been during a lecture at Sheffield that I first heard of William Rowan—the man who established the importance of change in day length in controlling reproduction and migration in animals. From that work in the early decades of the 20th century the science of photoperiodism grew, particularly in birds, the organisms Rowan had worked on. I also learnt that day that Rowan had succeeded under the most difficult of circumstances and that he had been badly treated in Canada where he worked.

Sixty-odd years on I have now read a biography of Rowan published in 1993.

The culprit who treated Rowan badly, even persisting with that animus when Rowan had achieved international fame for his research, was the founding president of the University of Alberta, Henry Marshall Tory (1864-1947), who took it on himself as a mathematician to decree the nature of proper zoological research as wholly laboratory based. He forbade fieldwork—an order Rowan ignored by working in his own time. Tory had been desperate to recruit somebody who could teach biology in 1919. A new medical building and an intake swelled by veterans of the First World War needed a zoology department since just as in Britain at the time, many medical students were unable to study biology at school and an elementary foundation, i.e. remedial, course mirrored that throughout the medical schools of the British Empire.

Henry Marshall Tory

Rowan’s biographer, the late Marianne Gosztonyi Ainley (1937-2008) skewers Tory’s reputation as an academic and research funding organiser and administrator, not only for his personal treatment of Rowan but also his short-sighted motives, his later leadership of what became the National Research Council of Canada and his disdain for the biology.

Born in Nova Scotia, Tory was a graduate of McGill University in maths and physics having begun his course at the age of 22. His mother was intent on her son becoming a Methodist minister and he then obtained a degree in theology, and spent two years in the church. In 1893 he jumped ship and became a lecturer in maths at McGill. To assist the setting up of a new physics department, Tory was sent to Cambridge for two terms to learn how things were done by the big boys. He thus became involved with two departments and worked his way up the academic ladder and doing administrative jobs for the university in setting up outstations, in British Columbia, for example, that in turn, became independent institutions.

In 1905, the new province of Alberta was formed and decided it needed its own university. The various accounts I have read state that Tory was appointed President in 1907. I found this rather odd since most universities in the Empire would have had either a Vice-Chancellor or Principal, following English or Scottish usage, respectively, for its chief executive position. McGill until recently had a Principal á la Glasgow and Edinburgh, before succumbing to the American and inaccurate usage of ‘President’ for the post.

There is no doubt that Tory was successful in building a new university from scratch. From his initial appointment of five academic staff in 1908 and admitting 32 students, Tory left in 1928 a university with five faculties, in modern buildings with 1,600 students.

Tory clearly regretted his appointment of Rowan almost as soon as he had made it. Indeed he went so far as to write to University College London to enquire if his degree was real, despite the fact that Rowan had continued to an MSc. The fact that Rowan did field work was anathema to Tory, which I suggest says more about Tory than it does Rowan who was equally at home with all aspects, approaches and techniques of zoology of the time.

Even when Tory had moved on to manage Canadian research funding  he continued to prevent Rowan from getting grants even though he had already obtained funding from American foundations and the Royal Society in London and his research was strongly supported by leading British zoologists of the time, Julian Huxley and James Peter Hill FRS (1873-1954) of UCL being two of them. Marianne Ainley wrote:

…Rowan submitted a major application to the NRC, but applied nowhere else. This was an error that he soon came to regret. After a decade of teaching and working under adverse conditions, Rowan must have realized that in Canada money for all but practical research was always scarce. He may not have known, however, that this was because Canadian science had been “guided by an entrepreneurial scientific ideology,” brought to this country by the original Scottish settlers.“ In the late 1920s, this “entrepreneurial ideology” still influenced the funding of Canadian science. Rowan often mentioned the lack of government money for basic biological research in his correspondence with Taverner, but, in his enthusiasm and naivety, Rowan paid no heed to the well-known fact that all the scientific departments of the federal government (the Geological Survey, Experimental Farms, and the Biological Board) had been established with practical aims in mind. In Canada, utilitarian science was supreme, and the NRC was no exception.

In early I930, Rowan joined the ranks of'Canadian scientists who continued to encounter discouragements and difficulties and even indifference to pure research. Tory’s presidency of the NRC further exacerbated the already difficult situation many scientists faced across the country, Tory was renowned as a vocal advocate of applied research, a firm believer in the usefulness of science. He was later described as a man “tended to favour the practical short-term problems that would make a noise; among the long-term projects, he favoured those with a staggering pay-off, preferably in tens of millions of dollars.” Tory’s attitude towards science exemplified the prevailing Canadian one. Unfortunately for Rowan, Tory was in a powerful position where he could prevent the funding of basic research and promote the projects of his choice, mostly those that involved applied research. A careful perusal of the list of projects funded by the NRC from I920 to 1935 shows, however, that some basic research was funded, particularly in Tory’s area, the physical sciences.

Tory also denigrated Rowan personally at this time even though it was Rowan who was putting Canada in general and the University of Alberta in particular on the map for world-famous and highly respected biological research. Ainley unearthed a letter written by Tory in 1932 and continued:

“During the last two or three years I was in Alberta I gave very little attention to Rowan due to the fact that only elementary work was done in the department” In fact, Tory’s last years in Edmonton (1925 to 1928) coincided with those of Rowan’s most intense research activity. And while Rowan carried out his early experiments during his spare time in his own backyard, nothing could long remain a secret in a small, closed, university community. Ironically, it was during this period that Rowan’s research on the effect of daylight on the reproductive organs of birds put the University of Alberta on the scientific world map. From Tory’s letter it is evident, however, that he continued to consider only laboratory work as real science, and chose to disregard Rowan’s pioneering investigations and subsequent fame. By ignoring Rowan’s ornithological research and the enthusiastic recognition given it by members of the larger scientific community, Tory could maintain that “only elementary work was done” in the university’s zoology department. For Tory, Rowan “had reached his limit” as both teacher and researcher. By taking this attitude,

Tory could with a clear conscience prevent Rowan from being funded by the NRC, and later destroy the younger.man’s chances for academic advancement.

But Tory did not finish there. He damned Rowan’s chances of getting the vacant chair of zoology at McGill. Despite glowing references from his supporters in London, Edinburgh and the USA, those responsible for the important took more notice of Tory who had been asked to provide an opinion. Ainley again:

…Tory replied: "With regard to Rowan, I find it a little difficult to write about him because I am afraid that what I have to say will not be of any great help to him in securing an appointment at McGill." Tory wrote, Rowan "got a B.Sc. from the University of London... on a semi-war basis," and informed Currie that he, Tory, had not been impressed with Rowan as a scientist, because Rowan "would not stick to the laboratory." Tory recalled he had told Rowan that "his only hope of becoming a competent zoologist and of proving his right to the headship of the department... would be by taking up some special line of work and sticking to it until he had proved his position as zoologist." Tory added, "Unless Rowan has completely changed since I knew him I would not consider him at all capable of ever organizing such a department or of drawing to it men who would be zealous for work.”

By contrast, Julian Huxley thought Rowan ‘a rare combination of fieldworker, systematist, general zoologist and experimentalist” who would make an excellent head of department.

I would argue that Ainley hit the elongated metal fastening on the cranium when she concluded that Tory was a product of the Scottish tradition. Here, I suggest, Ainley missed a trick. I suspect there were other influences at play. Tory clearly saw physics as superior in every way, as physicists so long have done, to the other pursuits that seek to explain and to exploit knowledge of the natural world. In McGill with Tory in the physics department was no other than Ernest Rutherford (1871-1937; later Lord Rutherford). It was Rutherford who is alleged to have said ‘all science is either physics or stamp collecting’. While there have been attempts to argue that Rutherford did not mean it quite like that, there is no doubt that if reflected a common view from the Age of Physics.

My view of Tory is perhaps summed by a line in Gilbert & Sullivan’s The Gondoliers, first performed in 1889.  The King of Barataria had become a  ‘Wesleyan Methodist of the most bigoted and persecuting type’.

With the tune of the Grand Inquisitor firmly fixed as an earworm, I will leave Tory and return to William Rowan himself and his biography by Marianne Gosztonyi Ainley to another article. But I least I know now how he was badly treated. 

Ainley MG. 1988. Rowan vs Tory: Conflicting views of scientific research in Canada, 1920-1935. Scientia Canadensis: Canadian Journal of the History of Science, Technology and Medicine 1, 3-21.

Ainley MG. 1993. Restless Energy. A Biography of William Rowan 1891-1957. Montreal: Véhicule Press.