Sunday 8 November 2015

The Frog and Parrot, or Dendrobates tinctorius and Amazona spp.

Dendrobates tinctorius - a form from French Guiana
No, not the name of a pub but a note about Dendrobates tinctorius, the Dyeing Poison-dart Frog from the Guianas and northern Brazil. The species was named ‘tinctorius’ by Cuvier in 1797 because of its use by Amerindians to change the colour of feathers of birds they kept, a process known as tapirage in French Guiana.

If you Google the scientific name, you will find links to a number of American zoos (often the purveyors of howlers and multiple but lesser degrees of misinformation) which describe the process as a legend foisted on early Europeans travelling in South America. However, as far as I can see the process seems to have a perfectly respectable provenance. The history of its description and information on where the process had been recorded was provided by Alfred Métraux* in 1928.

Dendrobates tinctorius - a yellow-backed form
The great Alfred Russell Wallace wrote the following in his book, A Narrative of Travels on the Amazon and Rio Negro, which was an account of his journey made from 1848 to 1852 and published in London in 1853:

They were all completely furnished with their feather ornaments, and I now saw for the first time the head-dress, or acangatara, which they value highly.This consists of a coronet of red and yellow feathers disposed in regular rows, and firmly attached to a strong woven or plaited band. The feathers are entirely from the shoulders of the great red macaw, but they are not those that the bird naturally possesses, for these Indians have a curious art by which they change the coIours of the feathers of many birds.They pluck out those they wish to paint, and in the fresh wound inoculate with the milky secretion from the skin of a small frog or toad. When the feathers grow again they are of a brilliant yellow or orange colour, without any mixture of blue or green, as in the natural state of the bird ; and on the new plumage being again plucked out, it is said always to come of the same colour without any fresh operation. The feathers are renewed but slowly, and it requires a great number of them to make a coronet, so we see the reason why the owner esteems it so highly, and only in the greatest necessity will part with it.

Dendrobates tinctorius
Hans Gadow (1855-1928) also described the process in his Cambridge Natural History volume, Amphibia and Reptilia, published in 1901:

It owes its specific name to the peculiar use made by man of the strongly poisonous secretion of the tiny glands of the otherwise smooth skin. Other species are doubtless employed in the same way. The poison is mainly used for “dyeing” the green Amazon-parrots. This is done as follows:—The green and blue feathers on the head and neck, or other parts, according to the fancy of the operator, are plucked out, and these places are rubbed with the poison, often simply with the living frog, certainly not with its blood, as is sometimes asserted. This operation may be repeated when the new, young feathers begin to bud. The result is that these appear yellow instead of green, and since the Brazilians, and to a certain extent the Portuguese, are rather partial to these artificially-produced freaks or “contrafeitos” as they call them, the industry is kept up.

There is doubt over which animals were used to paint the skin of the; Métraux, in his historical survey notes earlier writings on fish being used; there is also reference to other substances.

Dendrobates tinctorius, a form originally described
as D. azureus
However, back to the frog. Dendrobates tinctorius is highly variable, both in coloration and in body posture. Research using mitochondrial DNA and morphological characters firmly places all the different forms in one species, including the famous Blue Poison-dart frog, originally named Dendrobates azureus. The photographs shown here are of different forms of Dendrobates tinctorius that I kept and bred in my office in the 1980s and early 1990s.

Tapirage does raise questions that I have never seen answered. If the process changes the colour of newly-grown feathers to yellow instead of green, what is happening? Green is partly a structural colour in parrots made up of blue-green light selectively reflected by unpigmented ‘spongy cells’ which then passes through a yellow (i.e. blue-absorbing) pigment to leave green. With yellow the only colour showing after tapirage, there must be an absence of, or derangement in the spongy cells responsible for the blue-green, structural, element.

I have not found any reference to the examination of tapiraged feathers in such a manner that would provide the necessary information. Since photographs of ‘feather art’ are shown in articles, I would be surprised if a museum somewhere in the world did not have examples of tapirage.

The second question is what rôle, if any, is played by the skin secretion of Dendrobates tinctorius in tapirage? If important in achieving the colour change, are the toxins, mainly in this species, the pumiliotoxins, derived from the invertebrate eaten by the frogs, the chemical agents responsible?

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†Wollenberg K C, Vieth M, Noonan B P, Lötters S. 2006. Polymorphism versus species richness—Systematics of large Dendrobates from the Easter Guiana Shield (Amphibia: Dendrobatidae). Copeia 2003(4), 623-629

*Métraux A M. 1928. Une découverte biologique des Indiens de l’Amérique du Sud : la décoloration artificielle des plumes sur les oiseaux vivants , Journal de la Société des Américanistes 20 , 181-192. doi: 10.3406/jsa.1928.3646

‡Tinbergen J, Wilts B D, Stavenga D G. 2013. Spectral tuning of Amazon parrot feather coloration by psittacofulvin pigments and spongy structure. Journal of Experimental Biology 216, 4358-4364. doi: 10.1242/jeb.091561





Wednesday 28 October 2015

Galapagos Marine Iguana: Genes and Islands

The Galapagos never cease to fascinate. New discoveries are being made all the time on the animals that live there and how they have evolved. 

Marine Iguana. Santa Fé
(Barrington) 

I watched Marine Iguanas (Amblyrhynchus cristatus) for hours when we there in 2012: lekking behaviour, males fighting, mating, salt glands in action, feeding on algae, and just sitting around on rocks in large numbers.

Marine Iguanas vary in appearance between the islands and those with an obsession for artificial pigeonholes have created subspecies accordingly. I shall ignore those dated divisions and concentrate on a recent paper* in which mitochondrial and nuclear DNA sequences have been used to investigate the population genetic structure.

This paper has highlighted the tendency towards the splitting into different species—speciation—and the hybridization between lineages that tends to wipe out that tendency. While the split between the Galapagos Land Iguanas and the Marine Iguana was calculated to have occurred 4.5 million years ago, division into genetic lineages in the Marine Iguana was found to be very recent—within the last 50,000 years.

As well as differences between islands, different lineages were even found on one island: San Cristobal or Chatham as it was known in Darwin’s day, and to some of us still is. It is presumed that volcanic events, with their huge lava fields divided the populations at some stage; there was a major eruption 1800-3000 years ago which reduced the population and produced a severe bottleneck in both lineages. What is surprising is that migration and hybridization between these two populations, termed LO (for Lobería in the south-west of the island) and PP (for Punta Pitt in the north-east) was uncommon despite there being only 12 km of coastline between them. However, evidence was obtained for hybridization of both LO and PP lineages with animals from other islands rather than the adjacent population. Indeed, migrants were found from Santa Cruz (Indefatigable) and Espaniola (Hood) during the collection of samples. Therefore, introgression of genes from other populations acts to oppose the isolation that would tend towards speciation.


Marine iguana population clusters and phylogenetic relationships. (a) Map of the Galápagos archipelago with major islands colour-coded according to their marine iguana population cluster assignment inferred from structure analysis of 614 individuals genotyped for 12 microsatellite loci (vertical panel in (b)). (b) Species tree cloudogram based on an analysis of 6257 RADSeq-derived SNPs in 33 marine iguanas from across the archipelago, including both San Cristóbal lineages. The graph shows the posterior distribution of consensus trees. Asterisks mark nodes with posterior probability = 1.0 (all other nodes less than 0.9). Specimens were grouped according to population assignment based on structure analysis. From MacLeod et al. 2015


Genetic and morphological differentiation of LO and PP lineages on San Cristóbal Island. LO-SRL and PP-SRPC refer to the original Lobería and Punta Pitt localities, photos show adult LO and PP males. (a) Assignment of 454 individuals based on 18 microsatellite loci, after exclusion of inter-island hybrids and migrants. Abbreviations show sampling locations and 1993 marks specimens sampled in that year. (b) Haplotype network of control region sequences (mtDNA) for LO and PP specimens. (c) Map of sampling localities; arrows indicate migrants/hybrids from Santa Cruz (green), Española (orange) and Lobería (blue); dagger symbols denote locations of within-island hybrids between PP and LO; triangles denote locations of inter-island hybrids. Population SRECA contains Española migrants/hybrids only. Shaded areas mark lava groups 4–6 aged less than 0.1 Ma [40]. (c) Mean, standard deviation and range of morphological variables differing between LO and PP. ***p < 0.001, **p < 0.01, *p < 0.05; sample sizes above each plot, details and abbreviations in Results and electronic supplementary material. From MacLeod et al. 2015

There are differences in size between lineages, PP for example, being smaller than LO, but I will leave consideration of the significance of that and to what happens during El Niño oscillations, when food in the form of algae is short and the populations of Marine Iguana crash, to another time.

We only visited the south of Cristobal, so saw just those of the LO lineage. The question then, of course, is: which ones did Darwin see when he collected on Chatham (named, like Punta Pitt, after William Pitt the Elder, 1st Earl of Chatham, 1708-1778)? LO, certainly, in the south where there are groups of iguanas around Cerro Brujo, for example, but it is likely that he also saw PP, since some of his landing and collection points were well to the north and within the PP lineage’s range.

The photographs shows Marine Iguanas from some of the islands we visited:

Española (Hood)
South Plaza
Isabella (Albemarle)
Fernandina (Narborough)
The extent of the lava fields can be seen in this photograph taken on Isabella between from near Darwin Volcano to the south east:


A close-up of the lava on Santiago (James):



…and these are three videos, all taken on a wonderful morning on Fernandina (Narborough):



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*MacLeod A, Rodríguez A, Vences M, Orazco-terWengel P, García C, Reillmich F, Gentile G, Caccone A, Quezada G, Steinfartz S. Hybridization masks speciation in the evolutionary history of the Galapagos marine iguana. Proceedings of the Royal Society. B 282, 20150425.

†Estes G, Grant KT, Grant PR. 2000. Darwin in Galapagos: his footsteps through the archipelago. Notes and Records of the Royal Society of London 54, 343-368.

Friday 23 October 2015

Hedgehog Oil

Hedgehogs (Erinaceus europaeus) are said to be getting less common in Britain—hardly surprising news given the increase in the human population, the horizontal expansion of cities, towns and villages and the hard landscaping of gardens and public areas. The increase in the number of badgers which prey on hedgehogs is also being blamed.


The hedgehog population is these days though safe from organised human predation. I get bemused looks from my family when I tell them that the treatment in the 1940s and 50s for hard ear wax was hedgehog oil. From internet searches one may be left with the impression that hedgehog oil was used by country bumpkins who knew no better and who in turn had copied its use from the gypsies, as in this report from the Western Gazette of 11 March 1938:

Teddy Johnson, a 71 years-old recluse was on Sunday found dead in his small hut of earth and tins, near Swepstone, Ashby-de-la-Zouch, Leicestershire, where he had lived for 25 years. His past life was a secret he kept to himself. He sold watercress and an oil extracted from hedgehogs, which countryfolk regarded as a cure for deafness…

A Mrs O’Dell was singing its praises in Luton News and Bedfordshire Chronicle of 20 July 1950:

The oil from the hedgehog is the finest stuff in the world for earache. It has been worth a lot to me, with a family, as we could not get olive oil in war-time.
     My mother-in-law used to skin them, cook them slowly in the oven and strain off the oil. The hedgehog is a tasty dish.
     The gipsies like them and it was the gipsies who told us to use them for their oil.

Its use was, however, far more mainstream; hedgehog oil came in small bottles from regular pharmacies. The bottle and its contents would be warmed to body temperature in water or in front of the fire before the oil was run into the ear from an eye dropper. Olive oil and proprietary preparations gradually took over; and remember that was the only use olive oil had in the 1950s—if somebody had suggested cooking with it or even eating it with bread, they would have been seen as quite mad.

Searching for information on hedgehog oil is complicated by the fact that ‘Hedgehog’ was used as a brand name for lubricating oil by W.B. Harrison, a ship chandler, of Sunderland. 

The question is, of course, why was hedgehog oil so successful? It was liquid at room temperature and, unlike fish oil, had no unpleasant smell. The adipose tissue of domesticated animals is too hard at room temperature to be used. The melting point depends on the fatty acid composition and, yes, hedgehog fat has been analysed* and it is possible to see from the results why it can produce when rendered an oil at room temperature that is effective at softening cerumen and relieving the pain of hard wax.

…And you take this as a report from a satisfied user circa 1949.

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*Laukola, S. 1980. Seasonal changes in the fatty acid spectrum of the hedgehog’s white and brown adipose tissue. Anal. Zool Fennici 17, 191-201.

Tuesday 6 October 2015

The great fall of migrants on the Suffolk coast in September 1965; we were there

Male Bluethroat (Luscinia svecica)
photographed in Moscow
(from Wikipedia**)
Last month marked the 50th anniversary of a remarkable wildlife phenomenon on the Suffolk coast in the east of England. And we were there at the time.

As we approached the coast at Benacre on foot, we suddenly noticed that there were lots of birds around and that these birds were not the usual birds one might expect to see. In short, the shrubs and bushes were full of migrants and rare migrants at that. One bush had Bluethroats on nearly every branch and a Wryneck in the middle. There were Redstarts; Pied Flycatchers were abundant and a variety of small warblers skulked in the low shrubs and bushes. It was an amazing sight. Wherever we walked there were more and more birds.

A few days later when we returned they had all gone.

The ‘fall’ of migrants made the local weekly newspaper because Lowestoft, the town to the north, had been brought to a halt as exhausted migrants landed on buildings, roads and paths. in a post on Birdforum from 2008, a correspondent quoted from Birds and Weather: A Birdwatcher’s Guide by Stephen Moss (Hamlyn, 1995):

At just after two o'clock in the afternoon of 3rd September 1965, the residents of Lowestoft looked up to see a vast cloud of small birds overhead. Birds were dropping out of the clouds like raindrops, and soon the town was alive with them, in gardens, on the beach, and even in the roads, where many fell victim to traffic. Two people, in different parts of the town, actually had Redstarts alighting on their shoulders from the sky.
     Tens of thousands of birds were involved. All along the Suffolk and Norfolk coasts, great flocks of Northern Wheatears and Whinchats, Redstarts and Pied Flycatchers, Garden and Willow Warblers were arriving exhausted from the North Sea, and landing anywhere they could find food and shelter.
     The following morning, one observer, David Pearson, walked along the coast south of Walberswick. Along a 3-km stretch, he logged a staggering total of 15,000 Redstarts, 8,000 Northern Wheatears, 3,000 Garden Warblers, 1,500 Whinchats, 1,500 Tree Pipits, and 1,000 Willow Warblers, along with smaller numbers of other migrants. Rarer migrants were seen in unprecedented numbers, too: Wrynecks and Bluethroats reached double figures in several places, with Icterine and Barred Warblers, Ortolan Buntings and Red-backed Shrikes also appearing.
     The species involved and the time of year, left the lucky observers in no doubt that they were witnessing a massive displacement of Scandinavian migrants which, heading south-south-west across the North Sea to the coasts of mainland Europe, had been diverted westwards to East Anglia by the adverse weather conditions.

British Birds (59, 353-376) in 1966 carried an article by Peter Davis, then Migration Research Officer at the British Trust for Ornithology, entitled, ‘The great immigration of early September 1963’. It describes what was being seen by observers and ringers along the east coast of the British Isles over the few days in early September together with the weather maps. During the last few weeks of August there had been cloud and rain over Scandinavia. On 2 September there was an anticyclone over Scandinavia, creating ideal conditions for the birds to leave in very large numbers; at the same time there was a vigorous depression moving from France into the North Sea, bringing a north-easterly to easterly wind together with heavy rain on the northern edge of the depression, towards the Suffolk and Norfolk coast during 3 September. The migrants were pushed westerly and arrived in the massive falls recorded during the period of heaviest rainfall.

The fall of migrants was described as by far the heaviest of its kind ever recorded in Britain.

What we did not know when we were at Benacre was that a bird ringer was at work there. Davis reports:

…at Benacre Pits, most of the birds appeared between 13.20 and 13.50 GMT, many thousands arriving along a half-mile front. Pied Flycatchers and Redstarts were the most abundant, but there were huge numbers of Wheatears, hundreds of Whinchats, Garden Warblers and Willow Warblers, smaller numbers of Spotted Flycatchers, Tree Pipits, Whitethroats and Robins, and at least 20 Wrynecks. Between the start of the arrival on the 3rd and early on the 5th (when he ran out of rings) A. G. Hurrell caught some 650 migrants there, of which half were Pied Flycatchers (a much higher proportion than was seen at other places); also included were 14 Wrynecks, two Bluethroats, three Icterine Warblers and an Ortolan Bunting. 

Benacre Pits was part of the area we explored (probably on the afternoon of the 5th; we have no record). I think the Bluethroats must have avoided his nets since we saw many tens of individuals.

I should point out that Benacre has changed markedly through coastal erosion over 50 years. We walked down the same path to the beach a few years ago and found the chunk of land containing the bushes and scrub where we had seen most of the migrants had gone. We could not remember there being the cliff line then and I have just read in a blog that there was no cliff line here at all until 1981 ‘rather a gradually descending warren which over the years has been subsumed to the sea’.

Also gone were the Suffolk Punches, surely the most attractive of the draught horses, that lived in a field to the right of the path*.

This map from a modern Google Earth view was where we walked; it lacks the land that has disappeared into the North Sea between the end of the path and the beach. The point of the arrow shows where we first noticed that the bushes were full of birds.



Having seen the result of that massive fall of migrants, completely by chance, we have turned birdwatchers green with envy over a period of 50 years.

†Later Sir Anthony Hurrell, noted amateur ornithologist and bird ringer; a civil servant in the Ministry of Education for a time he was later British Ambassador to Nepal (1983-86). He died on 19 April 2009, aged 82.


*Were these the ones reported to have been used as two teams for ploughing, harrowing and towing balers on the Benacre Estate up to 1995? There area number of references to Suffolk Punches with Benacre in the name in studbooks and pedigrees.

**By Bogomolov.PL (Own work) [CC BY-SA 3.0 (http://creativecommons.org/licenses/by-sa/3.0)], via Wikimedia Commons

Friday 25 September 2015

How do hummingbirds drink nectar? And what about sunbirds?

The recent paper Hummingbird tongues are elastic micropumps published in Proceedings of the Royal Society has attracted attention in news media throughout the world.

The long-held notion that nectar flows by capillarity into grooves in the tongue that effectively form cylinders along part of its length seemed, first of all, to be supported by photographic studies and theoretical calculations. That paper was published, also in Proceedings of the Royal Society, in 2012 and entitled, The hummingbird’s tongue: a self-assembling capillary syphon.

Results in the most recent paper by Alejandro Rico-Guevara, Tai-Hsi Fan and Margaret Rubega were obtained by using high-speed cameras in eighteen species. It appears that the grooves, empty by compression inside the bill during protrusion and remain compressed until the tips of the tongue contact the nectar. The grooves then expand and fill completely with nectar. The tongue is withdrawn into the bill and the process is repeated. In other words, the observations are compatible with the mechanism being an elastic pump. With capillarity, these authors argued, the grooves should open to form cylinders before the tip of the tongue touches the nectar and a fluid meniscus should be apparent in the grooves. Those properties were not seen.


A video from the University of Connecticut (where two of the authors of the 2015 paper are based) can be seen here:


By contrast, the authors of the 2012 paper did report a meniscus in one species of hummingbird feeding from an artificial feeder, and the question is whether some difference in methodology or condition of the birds was responsible for the variation seen in the high-speed videos taken by the two groups. Two videos from that paper are shown here and here.

I suspect we have not heard the last of this story. It may not be one of those cases of either/or but of both, depending on circumstances as yet unknown. I can only add the observation that leaving things to capillarity does seem somewhat passive. In birds with such a high metabolic rate would it not be expected that an energy efficient pumping mechanism, faster than capillarity, would offer a selective advantage?

Finally, of course, the question must be asked: what is the mechanism in sunbirds, those unrelated, non-hovering nectarivores, of the Old World? Similar rates of nectar removal from flowers and artificial feeders to those recorded in hummingbirds have been reported*.

A female White-breasted Sunbird (Nectarinia talatala) that lived
in my office and at home at weekendsfor several years before
it went to join a male. It was fed exclusively on an artificial
nectar I designed specially for sunbirds

As a postscript it is worth noting that the titles of scientific papers are phrased very differently indeed. This is Gadow’s title for a paper on the subject that appeared in Proceedings of the Zoological Society of London in 1883 (pp 62-69): On the suctorial apparatus of the Tenuirostres.

It must be time to apply my suctorial apparatus to my dinner.

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†Kim W, Peaudecerf F, Baldwin MW, Bush JWM. 2012. The hummingbird’s tongue: a self-assembling capillary syphon. Proceedings of the Royal Society B 279 4990-4996.
‡Rico-Guevara A, Fan T-H, Rubega MA. 2015. Hummingbird tongues are elastic micropumps. Proceedings of the Royal Society B 282 20151014.
*Paton DC, Collins BG. 2006. Bills and tongues of nectar-feeding birds: A review of morphology, function and performance, with intercontinental comparisons. Australian Journal of Ecology 14 473-506.

Tuesday 22 September 2015

Found on a Hong Kong beach: What lifeforms are these?

AJP reports finding lots of these on the beach on the island of Po Toi last Sunday (a favourite place for birdwatchers since it is a major place for migrants to drop in) south east of Hong Kong island.



The discs are 3-4 cm in diameter.

After some searching he identified them as the remains of Porpita porpita, the Blue Button, superficially a jellyfish, but actually a colony of hydrozoan polyps, which is found in tropical and sub-tropical seas.

Now included in the Phylum Cnidaria, in our day they were in the Coelenterata along with the ctenophores.

Monday 21 September 2015

Hong Kong 70 Years Ago: Herklots the biologist; North the civil servant

The website Gwulo: Old Hong Kong is superb. Those who have followed the daily updates that show entries from the diaries written by internees in Hong Kong before, during and after the Japanese surrender, 70 years ago, will have noticed the name of Dr Geoffrey Alton Craig Herklots (1902-1986).

I hope to write more about Herklots later but at the time of the Japanese invasion in December 1941 he was Reader in Biology at the University of Hong Kong; he had joined the university in 1928 to start the teaching of biology. He later wrote the books, Hong Kong Birds and The Hong Kong Countryside. In addition to his job in the University, Herklots also became involved in Government work. In 1937, for example, he was appointed Superintendent of the Botanical and Forestry Department. He set up a Fisheries Research Station in the University and he founded, edited and contributed to The Hong Kong Naturalist.

Herklots was part of the group of underfed, ill and exhausted internees led by Franklin Gimson (who had himself sworn in as Acting Governor) out of Stanley to re-establish British rule immediately after the Japanese capitulation. The intense activity of those few days, before the arrival of the Royal Navy taskforce, is described in Philip Snow’s The Fall of Hong Kong (Yale University Press, 2003). Against a background in which U.S. army generals and Chiang Kai Shek’s Nationalists were trying to ensure that the latter took over Hong Kong, there were local deals to be done with the Japanese and with a local triad gang in order to maintain law and order as armed, looting gangs ran riot.


Camellia hongkongensis
from The Hong Kong Countryside
In the newly restored Government Herklots became Secretary for Development and had a particular responsibility for fisheries and agriculture. He became greatly involved in the business of survival both before and after the surrender to the Japanese in December 1941. Before the Japanese crossed the border, the Director of Medical Services, Selwyn Selwyn-Clarke, asked Herklots to devise a ‘siege biscuit’ that would provide the population with basic nutrition. With Thomas Edgar, a master baker, and after numerous trials, a successful product was launched. On the day before the Japanese attacked, a successful biscuit was produced that also contained calcium carbonate and shark liver oil.

In the Stanley internment camp, Herklots played closely involved with the welfare and education of its inhabitants who were malnourished, in terms both of quantity and quality, terrorised and short of medical supplies (even Red Cross parcels were held and not distributed by the Japanese). Herklots was a member of the organising Camp Temporary Committee, continued with running the University as far as was possible, gave general lectures ('Tropical Seas', for example) and continued work on trying to counter nutritional problems in the internees. This extract from Gwulo shows what he was involved in:

Thursday 1 July 1943
More than 18 months of poor nutrition are taking their toll. One of the most alarming developments is the occurrence of eye problems due to B vitamin deficiency. The camp's doctors are trying to tackle the problem, and today Dr Kenneth Uttley writes in his diary:
     Now that Lane Crawford’s baker Mr Edgar has been interned here, I have set Herklots and him on to the problem along with myself; we have cut down the amount of sugar required and are making more efficient use of the soya bean residue left after making the soya bean milk for the children and invalids. We have at last what appears to be a fairly active yeast and Geoffrey and I dispense it daily at 2pm to the eye cases and certain other B2 deficient cases.  Geoffrey and Edgar are busy most of each morning working on the yeast and are thoroughly enjoying themselves. We even entertain the idea of making yeast for the whole camp, but that will have to wait.
     Former Secretary to the Health Department J. I. Barnes reports that he was almost blind from vitamin deficiency but 'fully recovered' his sight after two doses of yeast - he got the second because he was a 'special worker' and most cases only got one dose. His job was looking after the camp stores - he slept there at night to prevent theft.

The internees in Stanley had also done a lot of thinking and planning. Before the arrival from London of a new civil administration to work under the temporary military administration, the ex-internees worked hard. At the speed that characterises the way things were done in Hong Kong, Herklots launched his new scheme for fish marketing on 12 September, less than a month after the Japanese capitulation, a week before the formal Japanese surrender and only 13 days after Harcourt’s ships and troops arrived in the harbour.

Herklots sketched his encounter with a large jellyfish
(The Hong Kong Countryside)
The other internee re-establishing British rule of interest here is Roland Arthur Charles North (1889-1961), Secretary for Chinese Affairs in the pre-war government and acting Colonial Secretary. I have written about him here*, since I have his complete run of The Hong Kong Naturalist on my bookshelves.

Herklots and North (a member of the old guard in the Colonial Service with whom Gimson disagreed on the way in which Hong Kong should be governed in the future) were members of the Hong Kong Government that seized the day and which issued its famous first communique since Christmas Day 1941 at 11.00 on 30 September 1945 The communique signalled loud and clear in thirteen words both immense relief and the carrying of a very big stick. Was it, I wonder, drafted by Duncan Sloss (1881-1964), temporary Director of Information and Vice-Chancellor of the University from 1937 until 1948?

Rear-Admiral Harcourt is lying outside Hong Kong with a very strong fleet…


The University of Hong Kong's building for biology in the 1930s
In the 1960s it housed the University Press. Before its conversion for biology  in 1928
it had been used for staff housing. After biology moved to the Northcote Building
it was briefly used as a women's hostel; Han Su-yin, under her university name of
Dr Elizabeth Tang was its sub-warden.
(from Bernard Mellor's The University of Hong Kong. An Informal History)



*I have made some recent additions to this post of 20 October 2013.

Wednesday 16 September 2015

Roger Akester (1922-2015) Avian Functional Anatomist

I was sad to see the death announced today in The Times of Roger Akester (Arthur Roger Akester), aged 92, on 29 August in Cambridge.

Roger Akester, veterinary anatomist at Cambridge from 1959 until 1991, will be remembered by all those with an interest in how birds work for his seminal contributions, made during the 1960s and obtained by x-ray and radio-opaque tracers, on the control of blood flow through the renal portal blood vessels and the passage of urine from the cloaca into the rectum and caeca by reverse peristalsis. The latter finding, which at the time was unexpected and contrary to the accepted view, provides the opportunity for modificaton of the salt and water content of the urine and for the uric acid to be metabolised by gut bacteria into useful products like volatile fatty acids; in other words, the recycling of waste products to get a second bite of the cherry.

The result of the mixing of urine, with its white uric acid, and faeces in the hind gut of birds is that both are voided together, as those who have been the target of a passing bird will testify. When I have to clean my car after an aerial assault by gulls that abound here, I cannot help thinking of both Erik Skadhauge, who went on the work on the salt and water aspects of modifying urine in the hind gut, and Roger Akester who discovered what was happening.

In the few hours since seeing the announcement in The Times, I have been able to find this radiograph (reproduced by W.J. Cliff in his book, Blood Vessels, Cambridge University Press, 1976), originally from Roger Akester's paper, on blood flow in the renal portal vessels and its control by the renal portal valves:


Tuesday 15 September 2015

Gustav Tornier and his tortoise, the Pancake Tortoise

Gustav Tornier
Mentioning what would now be regarded as completely crazy ideas of how a dirty environment induced the appearance of fancy goldfish in China and his hypothesis of 'plasma weakness' in my post on Dorothy Sladden and Ernest MacBride, I should note that Gustav Tornier (1859-1938) is best remembered for two things. The first was his incorrect representation of Diplodocus, a sauropod dinosaur, as a crawling, sprawled-leg reptile. The second is Tornier's Tortoise, now usually called the Pancake Tortoise, from rocky areas of East Africa, named after him by Siebenrock in 1903.

Siebenrock named the species Testudo tornieri; it is now Malacochersus tornieri. It was once commonly imported for the specialist reptile collector. Largely now protected, it is now commonly bred in captivity. It lays a single egg at a time—there isn't room for a clutch inside the flattened body.

This is the title page of Siebenrock's paper:



The start of the description of the new species:



The dedication to Tornier:


And the drawing:



And here is the nice short title of the Journal:



It was published in volume 112, pages 439-445.

We wrote about a then new finding in 1973 in an article in the Aquarist and Pondkeeper (38, 237-238):

The Pancake or Soft-shelled Tortoise...must be the oddest of all the land tortoises. The shell, which has the consistency of thick parchment, is thin and flat and entirely lacks a domed carapace. The bony plates which underlie the shell have large apertures so that the whole arrangement gives virtually no protection against predators. Early collectors found that these tortoises are extremely difficult to extricate from the crevices into which they run when disturbed, and suggested that not only do they wedge themselves in but actually inflate the body in order to press the shell against the walls of the crevice.
     In fact the ability to inflate was never checked although the story has been repeated from book to book since the 1920s with greater emphasis on the inflation than on the wedging action of the legs, even though Dr Robert Mertens had failed to find any evidence for inflation during the early 1940s. Drs L.C. Ireland and C. Gans of the State University of New York realised that the respiratory mechanism of tortoises probably would not allow the lungs to be inflated in order to distend the soft shell. Therefore they tested the response of Pancake Tortoises to being forcibly pulled from an artificial crevice they had been allowed to enter. Direct measurements of pressure in the lungs (reported in Animal Behaviour volume 20, pages 778-781, 1972) during this procedure clearly showed that there is no sustained inflation and that the tortoises depend on the wedging action of the fore- and hind-limbs to hold tightly in the crevice...

Later studies have suggested another mechanism. Richard Moll and Michael Klemens (Chelonian Conservation and Biology 1996 2, 26-35) looked at tortoises in the wild and reported that they could expand their bodies. They showed the presence of an extremely flexible diamond-shaped region on the plastron that protrudes when the legs are drawn in. This finding suggests that the lowered plastron region constitutes an important second type of wedge, and one that could easily have been interpreted as inflation of the body with air.

This is a photograph of captive Pancake Tortoises from Wikipedia. I have not been able to find one taken in the wild.


By Dave Pape (Own work) [Public domain], via Wikimedia Commons


That's him! Harold Munro Fox. A personal mystery solved fifty years later

Sometime early in 1965 I went for an interview for an overseas scholarship (overseas ones were funded by NATO) to work as a postgraduate student at the University of Hong Kong. The newly constituted Science Research Council was taking over from the Department of Scientific and Industrial Research as the British government's key funder of postgraduates. I remember very little about the interview, which was held at State House (demolished in the 1990s and after all but one Research Council were exiled to the outer darkness of Swindon) a then modern block in High Holborn in London, except for some very easy questions from an elderly man on the left. I also remember that I was not told who the three members of the panel were.

While looking up material for my jotting on Dorthy Sladden and Ernest W MacBride, I found myself looking at the photograph of the man on the left who dollied up the easy questions. It was Harold Munro Fox, FRS (1889-1967) best known for his work on animal pigments but also for his experiments which failed to replicate Kammerer's claims on regeneration in the sea squirt, Ciona.

Harold Munro Fox - from his Biographical Memoir
of the Royal Society

Saturday 12 September 2015

George Boyce (1920-95) of South Western Aquarists. Herpetologist

Some people have an influence far wider than their formal qualifications or occupation would suggest. In the world of herpetology in Britain of the 1950s and 60s, George Boyce, a pet-shop owner had a great influence on young people with an interest in reptiles and amphibians. Yes, he would sell you the animals but he was also a great enthusiast and advocate not only for reptiles but also for instilling a life’s interest in something worthwhile. This is a slightly modified version of this post on my other blog that deals with the history of the advances in keeping reptiles and amphibians.

In mid-1950s Britain dealers in reptiles and amphibians were rare for the simple reason that the market was small. Some larger dealers had gone out of business. Palmers of Camden Town was the really big one but did not specialise in reptiles. There were wholesale dealers selling to small pet shops but the main ones catering for enthusiasts and zoos were Robert Jackson in Cheshire and George Boyce in London. For a few years they were the only regular source.

George Boyce owned South Western Aquarists, a tiny shop at 2 Glenburnie Road, Tooting. It was a place where people, amateurs and professionals went to talk about and occasionally buy reptiles and amphibians. It was somewhat unusual because the reptiles and amphibians were not usually kept on the premises. From his lists which could be had by post, you could either get them delivered by rail to all parts of the country or you could write or phone to ask to look at something on the list; he would then bring the animals with him from home on the appointed day. Impulse buying was impossible. In the times I visited the shop over the years (1959 until 1982) I only ever saw a few lizards in one tank once. There were a few tanks of tropical fish and the then usual supplies of pet food. Conversations were frequently interrupted by the arrival of a little old lady to buy a pound of budgerigar seed or some dog or cat food. This local trade received the benefit of a level of attentive service verging on the obsequious before the owner returned to the subject of the conversation.

There is virtually no information on George Boyce or South Western Aquarists on the internet. However, I was delighted that he does make an appearance in the autobiography of Ken Livingstone, You Can’t Say That published in 2011 (London: Faber and Faber). For those readers not in UK I should explain that Ken Livingstone is a politician of the left and was Mayor of London from 2000 until 2008. He was a Member of Parliament from 1987 until 2001. He is also well-known as a keeper of reptiles and amphibians with the press usually referring disparagingly to ‘his newts’.

In describing his early life, he wrote:

Saturday afternoons were often spent at a pet shop on Tooting, chatting to the owner George Boyce about his imported reptiles, his Rotary Club activities and his Tory beliefs…

He goes on:

…Sadly, any chance of romance was dramatically reduced by my herpetological hobby. In all the Saturday afternoons I spent chatting to other reptile enthusiasts in George Boyce’s pet shop, only one young woman ever came in and she was dating another collector.

Ken Livingstone also reproduced a letter he had from George Boyce while backpacking to and from West Africa in 1966 which ends:

All the ‘lads’ and ‘lasses’ at the shop have asked me to convey to you their best wishes for your continued success in the termination of your journey. To these felicitations, Ken, I would add those of my own and of course Doris. Yours as ever, George.

I never went to Tooting on a Saturday on my rare visits to London. There was only ever George, me and his local customers. I remember him describing: visiting Jack Lester, Curator of Reptiles at London Zoo until he died in 1956, and Margaret Southwick, his assistant, on evenings when consignments of reptiles had arrived at the Zoo and were being unpacked; fishing at Frensham Ponds after the Second World War and catching large European Terrapins that had obviously been released there; the difficulties of dealing with exporters who would send what they had rather what was ordered; exporting imported animals to zoos in the U.S.A.

A frequent visitor to the shop and one who had just left when I got there was Dr Ian Wesley Whimster (1924-1979), Reader in Pathology at St Thomas’s Hospital Medical School. Part of his research was on the pigment cells in the skin of lizards; to that end he established a captive colony of Leopard Geckos.

He was a regular at meetings of the British Herpetological Society and wrote an article for the Journal on keeping crocodilians. He was there at the first meeting I attended (Summer 1961, I think) in the rooms of the Linnean Society in Burlington House. At that meeting everybody sat around a large mahogany table as live reptiles were passed around. I remember an impressive Leopard Tortoise and some superb Australian skinks owned by a woman sitting next to me. I asked how she had got so many and in such good condition. She explained that her husband was a pilot with B.O.A.C., the long-haul British airline that was merged with B.E.A. into British Airways, and he simply picked them off the taxiways and surrounding scrub at Sydney airport as he walked to his plane.

We also know from an article in Fishkeeping & Water Life that in 1958 he appeared on BBC Children’s Television describing Fish and How to Keep Them. In the BBC Genome project documenting BBC programmes the programme is there (20 January 1958) but he does not get a credit.

I have had some difficulty in finding more about George Boyce. However, thanks to his being in 1966 the executor of his father’s will (where he was described as aquarist and herpetologist) which was traced through a family history search website, I now have more information. George Frederick Boyce was born on 7 October 1920 in the Dorking Registration District of Surrey; in 1947 he married Doris L. Harrison in Croydon, Surrey. He died in 1995 in the Wandsworth Registration District of London.

I do not know when his shop closed; my guess is the mid- to late-1980s (my last visit to the shop was in November 1982). From advertisements in Water Life, it is clear that South Western Aquarists was operating in the late 1940s. Advertisements from 1950 mention a D.C. Crisp who must have been a partner, and Water Life (July-August 1950) in describing an aquarium show in London notes, D.C. Crisp of South London [sic] Aquarists made an intriguing picture, entering the hall with an alligator on a lead. Later in the 1950s, only the name of G.F. Boyce appeared on price-lists etc. and I have found no further clues on the role or identity of D.C. Crisp.

Here is an advertisement that appeared in Fishkeeping & Water Life in 1958:



Finally, a photograph of my Mississippi Alligator, photographed on the lawn of my grandmother’s second cousin’s house in Hendon, the afternoon I collected him from Tooting and eventually the subject of a note in British Journal of Herpetology*.


As always, any further information on George Boyce is welcome. Please contact me if you have any.


*Peaker, M (1969) Active acquisition of stomach stones in a specimen of Alligator mississippiensis Daudin. British Journal of Herpetology 4, 103‑104