Citreoline Trogon: a bird endemic to western Mexico

Citreoline Trogon
1 February 2025, El Jorullo, Jalisco, Mexico

We saw this Citreoline Trogon after walking across the 470 metre long suspension bridge at El Jorullo a few miles inland from Puerto Vallarta on the Pacific Coast of Mexico. We had to wait for a while because Canopy River park is used for all sorts of outdoor activities and several convoys of all-terrain vehicles were heading in the opposite direction. The suitability for such vehicles makes it, so it is claimed, the longest vehicular suspension bridge in the world. The whole area has been owned and operated since 1940 by an association of local foresters; the vehicles and zip wire etc came later, after 2005. The Sierra Madre mountains provide a backdrop to the protected area of forest while the Cuale River runs way below the bridge.

Jorullo Suspension Bridge

The Citreoline Trogon (Trogon citreolus) is endemic to the woods and forests of western Mexico. They nest by digging into the nests of tree termites. This one was eating berries and at various times of year eat either or both fruit and insects.

From the photograph at high magnification it can be seen that the trogon has two toes visible at the front. Does that make it a zygodactyl like parrots and woodpeckers? Well, no. It does have two toes at the front and two at the back but in trogons digits 3 and 4 face forward while 1 and 2 face back. In zygodactyly, 2 and 3 face forward and 1 and 4 back. Trogons are thus described as heterodactylous.

There were mammalian delights too along the trail through the forest beyond the suspension bridge but they are for later.

Comparative Endocrinology Revisited: Hellbenders, Giant Salamanders and their hormones

Hellbender from the Appalachian Mountains
Brian Gratwicke Creative Commons Attribution 2.0 Generic

In a recent article I mentioned that there is still discussion on why the adrenal glands of different groups and species of vertebrates produce either or both of the two steroid hormones, cortisol and corticosterone. Both are these hormones are glucocorticoids, originally named for their effect on glucose metabolism and both have wide-ranging effects on the body. Both have a little mineralocorticoid action, i.e. acting to retain sodium, the role of the other major adrenal steroid hormone, aldosterone. In different tissues of different animals corticosterone and cortisol have been found to have slight differences in their mineralocorticoid action.

The difference between corticosterone and cortisol is the absence or presence of a hydroxyl moiety at the 17 carbon position of the steroid molecule. Corticosterone does not have it; cortisol does. In some mammals, rats and mice for example, the adrenal cortex does not have the enzyme 17α-hydroxylase* An earlier chemical precursor of cortisol is therefore not produced, leaving corticosterone as the only glucocorticoid produced by the rat adrenal. Since aldosterone has been found in all vertebrates, corticosterone must also be produced since it is a precursor in aldosterone synthesis. In some animals that produce both corticosterone and cortisol, one or the other may predominate. In primates it is cortisol.

The structural difference between corticosterone and cortisol (ringed in red)
Adapted from the classic textbook by Aubrey Gorbman and Howard Bern,
A Textbook of Comparative Endocrinology, New York: John Wiley. 1962

A paper published in 2020 raised an interesting question and demolished the generally accepted view that corticosterone is the major and perhaps the only major glucocorticoid produced by the adrenal cortex of amphibians. Hellbenders (Cryptobranchus alleganiensis) are the American relatives of the Giant Salamanders of East Asia. They too are large, but not so large, up to 70 cm in length, long-lived and fully aquatic. As part of an extensive study of an amphibian in decline in the wild, the concentration of cortisol was found to be five times that of corticosterone in blood taken from a tail vein. The difference was even greater—up to ten times—after the injection of mammalian adrenocorticotrophic hormone (ACTH). The authors suggested that cortisol may predominate, as it does in fish, in these aquatic salamanders since in other vertebrates cortisol has a somewhat greater mineralocorticoid activity than corticosterone thereby preserving salt and water balance. However, that inference, at first sight, does not seem to hold. In work done in Hong Kong by my old fellow postgraduate student Samuel Chan, Tom Sandor (1924-2003) (a doyen of steroid biosynthesis in vertebrates) and Brian Lofts (1929-2015) only corticosterone was found after adrenal tissue from similarly aquatic Chinese giant salamanders (Andrias) was incubated with radiolabelled precursors in the steroidogenic pathway.

The authors of the papers on the Hellbender drew up a table showing previous studies on the occurrence of corticosterone and cortisol in amphibians and emphasised the difficulties of interpreting studies in which blood concentrations were recorded and experiments on adrenal tissue in vitro. They also noted that a number of studies were done before reliable methods had been established for identifying and quantifying corticosterone in the presence of cortisol and vice versa. They made sure that they established the chemical identity of the hormones using mass spectroscopy—something not possible in the early studies with the low but biologically active concentrations found in blood plasma. However, there is another difference that may be significant: the site from which blood was taken. Because the analytical methods did not exist at the time to study hormones in peripheral blood, John Phillips (1933-1987) made his name in the late 1950s by devising techniques to sample blood from the veins draining the adrenal gland or from those that carried a high proportion of adrenal venous effluent. The distinction is important because, as I suggest below, any production of adrenocortical-like hormones from elsewhere in the body would not have been at detectable concentrations.

What could be the explanation of the findings in the Hellbender and indeed in those amphibians in which both corticosterone and cortisol have been found to be present in the blood (albeit with the caveats on methodology outlined above)? I suggest there are two hypotheses which are not mutually exclusive:

1. The adrenal tissue expresses the gene CYP17A1 that produces 17α-hydroxylase during the breeding season but not at other times. The Hellbenders sampled were, as the authors note, caught in late summer ‘prior to nesting because at this time adult male and female hellbenders can be distinguished by the swollen cloaca of males’. There is evidence gathered in the first half of the twentieth century that the adrenal of many amphibians varies in appearance, both macroscopic and microscopic, according to season. In newts and salamanders the adrenals are multi-segmented strands of tissue on the ventral side of the kidneys. In the California Newt (Taricha torosa), as one example, during the two-month breeding season the adrenal is bright orange in colour and expanded compared wit the non-breeding season when the gland is dull, smaller and flatter. In other words, the gland waxes and wanes and it is easy to imagine that the production of 17α-hydroxylase follows a similar pattern. An adrenal with 17α-hydroxylase is also capable of producing androgens that might play a role in reproduction.
2. The adrenal is not the site of cortisol synthesis or secretion. The gonads in a number of vertebrates are known to be capable of synthesis and secretion of adrenocorticosteroids. Of necessity in being able to sythesise androgens and oestrogens they must express CYP17A1. The adrenal, testis and ovary are all of similar embryonic origin and complete separation of steroid synthesis and secretion   is far from complete with overt clumps of cells characteristic of the adrenal cortex being present in the ovary.

A possible clue as to the cellular origins of of the two adrenal steroids is that while in the Hellbender both corticosterone and cortisol concentrations increased in response to ACTH the time course was different and there was no correlation between concentrations of the two hormones in blood at any time point. Thus corticosterone reached a peak at one hour after ACTH; cortisol at three hours. Does this finding suggest a different cellular origin of the two steroids? I suspect that it does with either two cell populations in the adrenal itself (one only seasonally present producing cortisol and probably androgens, the ‘permanent’ cells producing corticosterone) or of the adrenal producing corticosterone and the ovary and testis producing cortisol as well as androgens and oestrogens.

Assuming that the other members of the cryptobranchid family have a similar endocrine system, both of these possibilities are compatible with the finding of cortisol and corticosterone in the blood of Hellbenders but the finding of only corticosterone  being produced in vitro by the adrenal of Chinese giant salamanders, assuming of course the latter were not in the breeding season or breeding condition.

Chinese Giant Salamander (Andrias sp)
On the roof of the now demolished Northcote Science Building
Univewrsity of Hong Kong ca 1966

I would argue that the next step in solving this problem lies in China where farm-bred Giant Salamanders are readily available in the restaurant trade. The question is a simple one: what are the concentrations of corticosterone and cortisol in the blood at different stages of the reproductive cycle? Modern students may prefer the question in molecular biological terms: where are the sites of expression of the gene CYP17A1?

...And surely the Axolotl is just asking to be looked at!

Comparative endocrinology may no longer be fashionable but there are still important areas where new knowledge and a look at unsolved old problems are needed if we are to understand how signalling systems that operate between organs and tissues work. The science of intercellular signalling is now being neglected. 

*I have seen the odd paper showing cortisol as present in mice (at least in a particular strain of lab mouse) but without comment as to the significance of the finding.

The paper on the Hellbender:

Hopkins WA, DuRant SE, Beck ML, Ray WK, Helm RF, Romero LM. 2020. Cortisol is the predominant glucocorticoid in the giant paedomorphic hellbender salamander (Cryptobranchus alleganiensis). General and Comparative Endocrinology 285, 113267 doi: 10.1016/j.ygcen.2019.113267

The paper on the Chinese Giant Salamander:

Chan STH, Sandor T, Lofts B, 1975. A histological, histochemical, and biochemical study of the adrenal tissue of the Chinese giant salamander (Andrias davidianus Blanchard). General and Comparative Endocrinology 25, 509–516 doi: 10.1016/0016-6480(75)90161-6

For the older literature on the adrenal gland of amphibians:

Chester Jones I. 1957. The Adrenal Cortex. Cambridge: Cambridge University Press.

UPDATED 17 JUNE 2025

Crocodiles and Birds at the Mirador de las Aves near San Blas in Mexico

We have seen lots of crocodilians in the wild but rarely doing anything but sunning themselves or just lying in the water. When we pulled into a lay-by complete with food stalls and an iron-railinged viewing area on the outskirts of San Blas in Mexico to look at the birds on the lagoon and in the surrounding mangroves, I had read that American Crocodiles (Crocodylus acutus) could be seen there. The lay-by whose name gives a clue as to its avian delights, Mirador de las Aves, is clearly a popular place to stop, buy a snack and look at the wildlife as locals drive along the road on a Sunday evening.

And there were crocodiles, large ones hauled out just under the viewing platform and basking in the sun on the edge of the lagoon. However, as the sun went down and moved off the crocodiles they entered the water and several apparently began searching the bottom for food with one, as the video below shows, emerging to swallow something small it had caught or found. Eventually I counted seven moving around offshore or simply hanging in the water as dusk fell.

Animals of the Namib Desert. The South African Zoologist Who Worked Out How They Survive On the Foggy Foggy Dew

Last month we had a very brief visit, counted in minutes not hours, to Stellenbosch in South Africa. A few days later we were visiting Namibia for a couple of days and for a second time. The Namibian desert with its sand dunes is famous for the ways in which the various species of animals and plants that live there obtain water. The Namib is very different to what many think of as a desert because although it lacks rain it gets regular and small amounts of water delivered to it in the air.  In short it is a land of fog and dew. The sea off Namibia is cold by virtue of the Benguela current running north. Thus fog banks form along the coast which are blown into the western part of the Namib. Water vapour condenses on surfaces during the cool night and penetrates the soil during the hot day. We have all seen on television the ways in which some beetles channel condensed water collected on their bodies into their mouths.  Snakes lick condensed water collected on their scales. Dune lizards, which occur only in the fog belt of the Namib, by contrast, drink avidly from surfaces and sand but then store water in the expanded caecum for several months. The Hairy-footed Gerbil (Gerbillurus paeba), unlike some other desert rodents, cannot not survive on dried seed alone; it relies on water from the succulent plants that in turn rely on the fog and dew.

The only university building we had time to see in Stellenbosch
housed the Faculty of Theology

Namibia and Stellenbosch were connected in my mind because exactly 50 years ago we had a visiting worker in the department, the man who put the desert animals of Namibia on the map. Gideon Louw* spent January to June 1975 working on mammalian thermoregulation with John Bligh. Although the rest of the department did not see much of him, he did come along for a chat several times and we discussed all sorts of things about how all sorts of animals work in all sorts of environments. I remember the impression of a very nice man but deeply troubled by events in his own country. He was, I now learn, then shortly to move from his position as Professor of Zoology in Stellenbosch to the University of Cape Town. Biographers have recorded that Gideon Louw was a founder member of the Progressive Party which was utterly opposed to the apartheid policies being imposed by the government then in power. Stellenbosch was I have also learnt a  stronghold of support for the National Party and an uncomfortable place for those opposed to its policies.

Gideon Louw

In an appreciation of the work of Gideon Louw published in 2004, Joh Henschel and Mary Sealy wrote (references  omitted):

In hyperarid areas such as the Namib Desert the occurrence of fog and dew plays an important role in the water economy of many desert organisms. This fact was known, particularly through the works of Walter (1936) and Koch (1961), by the time that Louw first visited the Gobabeb Training & Research Centre in the Namib Desert in 1966.

Louw soon recognised that ecophysiological mechanisms were a key towards understanding the relationship between atmospheric moisture and desert organisms. Extensive contributions of this doyen of desert ecophysiology range from water, energy, and salt balance, to thermal biology, conjunctively in terms of physiology, behaviour and ecology. He inspired a generation of students and colleagues to elucidate these mechanisms and their consequences, and later reviewed some of these insights.

Like a number of others working in his field, Louw did not have a first or even a higher degree in zoology. He had started on the animal science of agriculture and his first job was as an agricultural geneticist. Several spells in the USA, including a PhD in endocrinology at Cornell, followed before he returned to South Africa and a personal chair in zoology at Stellenbosch.

Louw’s work in Namibia became well known internationally when he was invited by Geoffrey Maloiy, the organiser, to a Zoological Society of London Symposium ‘Comparative Physiology of Desert Animals’ in July 1971, a meeting I had to miss because of a clash of dates. The published volume is still worth reading since all the major players in the field were contributors and showed the state of play after a major burst of activity that followed World War II.

Gideon Louw did much more than his research on the animals and environment of the Namib including writing several books, while his students have spread across the world and are well-established names in the biological sciences. From the reading I have done it appears that he was appalled by the political violence sweeping South Africa in the early 1990s and retired to Canada. He died on Vancouver Island on 22 March 2004, aged 73.

SJP spotted this lizard in the dunes at the deserted mining village of Kolmanskop
near Luderitz in Namibia. It is the Common Rough-scaled Lizard(Meroles
squamulosa) a psammophilous species closely related to the dune lizard
(M. anchietae) studied by Gideon Louw
(Noble Caledonia Expedition Team Photograph)

This beetle was on the dunes at Kolmanskop

Lichens on rocks in the desert get their water from the dew

The reason this lichen is so green is that there had been rain
sufficient for water from the Swakop River to empty into
the Atlantic as in 2011 and 2022

A wide-angle view from the 'moonscape' viewpoint inland from Swakopmund

*Gideon Nel Louw, born 24 December 1930 East London, South Africa; died 22 March 2004.

Anon. 2004. Tribute to world-renowned zoologist. University of Cape Town News HERE

Cherry M. 2005. Gideon Louw (1930-2004). South African Journal of Science 101, 399.

Henschel JR, Seely MK. 2004. First Approximation of the Ecophysiology of Fog and Dew – A Tribute to Gideon Louw. Proceedings of the Third International Conference on Fog, Fog Collection and Dew: Cape Town, South Africa, 11-15 October 2004 [no page numbers shown].

Louw GN. 1972. The role of advective fog in the water economy of certain Namib desert animals. Symposia of the Zoological Society of London 31, 297-314.

Roseate Spoonbills Feeding in Mexico

 

Roseate Spoonbills (Platalea ajaja) were feeding in a roadside lagoon near San Blas on the Pacific Coast of Mexico in early February. Having seen all six species of spoonbill on our travels around the world these were by far the most active we had come across. The frantic sweeping of the bill left and right, the sudden snap shut and the lift to swallow whatever they have caught is an amazing spectacle. If you look closely at my video you will see one catches and swallows a small fish.

There have been all sorts of suggestions as to how spoonbills detect the prey passing between the upper and lower ‘spoons’ and thus rigger the bill to snap closed. There is a good discussion of the topic by Bruce Taggart HERE.

We passed this lagoon (with a restaurant by the edge) several times. Each time it had large numbers of Great (Great White) (Ardea alba) and Snowy Egrets (Egretta thula), as well as the spoonbills.

Do Not Annoy an Amphiuma

I came across a video on YouTube which reminded me of what happened 49 years ago and the tale that followed. In previous articles I have recounted the appearance of the now critically endangered giant salamanders of China in the zoology department of the University of Hong Kong for class dissection. We took some on to the roof to photograph them before they were killed ready for the students  One lunged with its mouth open and snapped at SJP’s hand which she withdrew rather rapidly thus avoiding a bite.

Back down in the lab we were told that John Phillips, the head of department, had been bitten badly by another amphibian, an amphiuma, and that he had, as a result, had a recurring fever which took a long time to go away.

I should say that I have never seen an amphiuma, alive or dead, but to anglers and commercial fishermen in the right part of the USA they are a familiar sight. Amphiumas are eel-like salamanders which can grow to over a  metre in length. They are virtually entirely aquatic, living in swamps, lakes and ponds, but will move on land after heavy rain. They are nocturnal and carnivorous.

What was John doing with amphiumas I asked myself, knowing that in the late 1950s or early 1960s it would be something to do with the hormones produced by the adrenal cortex. After a bit of searching I found a reference that provided some of the information. The results were reported as part of a paper given at a symposium on comparative endocrinology held at Cold Spring Harbor in  May 1958. John had arrived in the USA in September 1957 after working for his PhD in Liverpool. I cannot be certain whether his encounter with an amphiuma was in Liverpool or shortly after his arrival at Yale on Fellowship of the Commonwealth Fund of New York. I cannot get hold of the paper (published in 1959) but I suspect it did not give much further information on where the various studies were done. Since he is known to have continued his work on adrenocortical secretions in vertebrates at that time, my bet would be Yale. Furthermore, amphiumas were readily available in the USA from biological suppliers.

The amphiuma in question was a Three-toed Amphiuma (Amphiuma tridactylum) easily distinguished by the number of toes on its tiny legs. Its distribution is ‘centered on the lower Mississippi River from Texas to western Alabama, and north to southern Illinois and extreme southwestern Kentucky’. They were and probably still are a source for laboratory suppliers since they are a bycatch from crayfish trapping.

Three-toed Amphiuma
Photo by Ashley Wahlberg (Tubbs) via Flickr

All descriptions of amphiumas note that they bite savagely as well as being very difficult to handle because of the slimy mucus on the tough skin. It is no wonder amphiumas are called ‘eels’ as in ‘congo eels’, a term derived from conger eels. No wonder John Phillips got bitten and became infected by some pathogen lurking at the bottom of a swamp.

As the following video shows, it is not only when handling amphiumas that they are liable to bite.

Seventy years since John Phillips started his pioneering work there is still active research and discussion on why the adrenal glands of different groups and species of vertebrates produce either or both of the two steroid hormones, cortisol and corticosterone. Indeed, the field is just hotting up again.

*John Guest Phillips FRS.1933-1987.

Chester Jones I, Phillips JG, Holmes WN. 1959. Comparative physiology of the adrenal cortex. In Comparative Endocrinology: Proceedings of the Columbia University Symposium held at Cold Spring Harbor 1958. (ed. A. Gorbman), 582-612. New York: Wiley.

Sandor T. 1969. A comparative survey of steroids and steroidogenic pathways throughout the vertebrates. General and Comparative Endocrinology Supplement 2, 284-298.

Vinson GP, Tait JF. 1988. John Guest Phillips. 13 June 1933-14 March 1987. Biographical Memoirs  of Fellows of the Royal Society 34, 610-637 doi: 10.1098/rsbm.1988.0020

Identification of the Coelacanth in 1939. A fishy story or just a fishy story?

If the 1950s had a celebrity species, it was the Coelacanth. Mysterious and only recently discovered to be alive rather than existing as fossilised remains, there was massive media coverage of this ‘living fossil’. The second specimen was discovered in 1952 and the story was kept fresh by the publication of J.L.B. Smith’s book published in London as Old Fourlegs. The Story of the Coelacanth and in New York as The Search Beneath the Sea – The Story of the Coelacanth, in 1956.

Coverage of the story was all over the media and I remember late one night in the late 1950s staying at my grandmother cousin’s house in Hendon, north London, searching for something to read in her extensive bookshelves. I picked up a copy of Reader’s Digest and there was the intriguing title of an article, ‘The fish named L.C. Smith’.  The title was of course an abbreviation of the scientific name, Latimeria chalumnae Smith 1939.

Rhodes University

The central character of the coelacanth story is James Leonard Brierley Smith (1897–1968). He was an organic chemist, educated in South Africa. He then did research in Cambridge. He received the 14th Ph.D. degree to be awarded by that university in 1922 for a thesis entitled, Interaction of sulphur monochloride with substituted ethylenes. He was plagued by illness which began while serving in the army during the First World War; this continued after he had returned to South Africa and life as a lecturer in the chemistry department of the University of Grahamstown. An interest in sea fishing developed into research in ichthyology an arrangement that often did not meet with the approval of the university. He was, when it came to fish, a professional scientist and university lecturer but an amateur ichthyologist. By the late 1930s he had become though the acknowledged expert on the fishes of South Africa.

The basic story of the discovery of the first specimen is well known.  On 22 December 1938, Marjorie Courtenay-Latimer (1907– 2004), the enthusiastic curator of the East London Museum in Eastern Cape, answered a telephone call  from the skipper of a trawler asking if she would have a look at some interesting bycatch. One fish stood out: five-feet long, beautifully coloured with hard scales, limb-like fins and a distinctive tail. She did not know what it was. She and an assistant carried it in a taxi to the museum but after consulting her books still could not identify it. She then wrote to J.L.B. Smith at Rhodes University in Grahamstown including in her letter a drawing of the fish. But he was away and Miss Courtenay-Latimer grew increasingly desperate as the fish (refused entry into the local mortuary’s frig and the cold store) by now wrapped in formalin-soaked blankets in the premises of the town taxidermist was starting to go off. Eventually, and desperate to preserve something, she had the taxidermist skin and stuff the fish in the manner of a ‘game’ fish. Smith’s eventually received the letter while he was still away from Grahamstown in Knysna, 350 miles from East London. He immediately realised from the drawing that he was looking at something special when it dawned on him that it could be a coelacanth. Scales from the fish were sent to him which convinced him that it was indeed a coelacanth. When he and his wife were back in Grahamstown they travelled to East London to see the fish. It really was a coelacanth. And the world went coelacanth crazy as the news got out. The fish had been caught off the mouth of the Chalumna River. Smith therefore came up with the scientific name to mark the achievement of Marjorie Courtenay-Latimer in realising the fish was something special and doing everything she could to get it preserved and identified together its origin. One might have thought that commemorating the Captain of the trawler, Nerine, Handrik Goosen, might have been more appropriate since he could easily have chucked the by-catch coelacanth over the side; instead he consulted the local museum.

from Smith 1956

Key locations in the identification of the coelacanth
Having passed down this coast by ship last month I wonder how close
we came to a living coelacanth

from Smith 1956

The coelacanth from December 1938 as Smith saw it on reaching East London
from Smith 1956

But there is a twist. While looking for something else entirely I came across an excellent history of the entomology and zoology department of Rhodes University in Grahamstown (now Makhanda) and found this:

[J.C.] Van Hille always considered that Alice Lyle never received credit for her achievements. He also believed that she played a critical role in the identification of the first coelacanth (Latimeria chalumnae) in 1938. In a letter to Miles dated 30 April 1980 van Hille wrote: “The zoo lecturer you refer to was Alice Lyle, she was a dear! It was actually she who identified the coela canth. Miss Latimer, of the East London museum saw that it was something quaint and kept it in formalin in her bath. She sent a little sketch to J.L.B. [Smith] who was on holiday in Knysna and was slow in reacting. Eventually he took the sketch to Omer-Cooper {the head of department and an entomologist] who also had no idea. Alice came in to bring them tea and saw it and said ‘I have seen it before’. She studied in Bloemfontein [Grey University College which became the University College of the Orange Free State] where the emphasis was on vertebrates. Omer was more of an Entomologist and Smith a Chemist. So Alice got the Cambridge Natural History from the library and there was a good picture of the coelacanth. I [van Hille] have this story from Omer and always think of it when in all the books and articles it mentions that J.L.B identified the coelacanth.” Lyle resigned from Rhodes in 1949 for an appointment as Guide Lecturer at the Durban Museum, where she also lectured to 1st year medical students at Natal University. She was appointed to Fort Hare as Professor of Zoology in 1952 but sadly died in October of that year in Cape Town.

I have found no indication that it was Alice Lyle who identified the drawing as a coelacanth before Smith. His letters, reproduced in his book, show that he was virtually convinced it was a coelacanth before leaving Knysna and had written to Courtenay-Latimer to say so. It is tempting to speculate that Smith may have gone to the zoology department looking for illustrations of coelacanths and that Alice Lyle found one in the Cambridge Natural History Volume 7. However, that illustration is from Arthur Smith Woodward’s 1888 catalogue of fossil fishes in the Natural History Museum in London and we know from Smith’s book that he had already consulted that volume, having had it sent from Cape Town to Knysna when he realised the possibility that he may be dealing with a coelacanth. Indeed, he adapted the illustration from Smith Woodward for his 1956 book.

Illustration in the Cambridge Natural History

from Smith 1956

Smith’s visit to the zoology department must have been during the second week of February 1939. As shown in the quoted passage below, Smith had intended to drive straight from Knysna to East London. However, the roads were difficult because of heavy rain and he had to head for Grahamstown and stay there a week before reaching East London on 16 February.

We are left pondering. Is the story of Alice Lyle identifying the coelacanth, in fishy terms, a red herring (zoologically, a clupea rubra)? Was a conversation misheard in which Smith consulted colleagues to see if they agreed with his tentative identification? Beyond that sort of explanation we would be in the realms of conspiracy theory.

Finally, the passage from Smith’s book in which he saw at long last what became the most famous fish in the world:

We left Knysna on the 8th February 1939, intending to go straight through to East London, but there was to be nothing easy about this, for we travelled in continuous heavy rain and were fortunate to reach Grahamstown, since by that time floods had rendered almost all roads impassable. Drifts and slippery mud made motoring in South Africa no light undertaking in those days. We had to wait a whole week before the roads to East London became usable, and after an awful journey reached there on the16th February 1939.

We went straight to the Museum. Miss Latimer was out for the moment, the caretaker ushered us into the inner room and there was the—Coelacanth, yes, God! Although I had come prepared, that first sight hit me like a white-hot blast and made me feel shaky and queer, my body tingled. I stood as if stricken to stone. Yes, there was not a shadow of doubt, scale by scale, bone by bone, fin by fin, it was a true Coelacanth. It could have been one of those creatures of 200 million years ago come alive again. I forgot everything else and just looked and looked, and then almost fearfully went close up and touched and stroked, while my wife watched in silence. Miss Latimer came in and greeted us warmly. It was only then that speech came back, the exact words I have forgotten, but it was to tell them that it was true, it was really true, it was unquestionably a Coelacanth. Not even I could doubt any more.

Live coelacanth seen off Pumula on the KwaZulu-Natal South Coast, South Africa, 2019
Bruce A.S. Henderson, CC BY 4.0
<https://creativecommons.org/licenses/by/4.0>, via Wikimedia Commons

Go! & Express. 27 February 2020
The coin was minted in 1998

Bruton, MN, Coutouvidis SE, Pote J. 1991 Bibliography of the living coelacanth Latimeria chalumnae, with comments on publication trends. Environmental Biology of Fishes 32, 403-433.

Dugan J. 1955. The fished named  L.c. Smith. Reader’s Digest, December 1955, 147-151.

Hodgson AN, Craig AJFK. 2005. A century of zoology and entomology at Rhodes University, 1905 to 2005. Transactions of the Royal Society of South Africa 60, 1-18.

Smith JLB. 1956. Old Fourlegs. The Story of the Coelacanth. London: Longmans, Green and The Search Beneath the Sea. The Story of the Coelacanth. New York: Henry Holt.

Dr Lyon in 1916 questions: “Who the hell put the ‘l’ in Ambystoma” - or words to that effect

Spotted Salamander, Ambystoma maculatum
Peter Paplanus from St. Louis, Missouri, CC BY 2.0 <https://creativecommons.org/licenses/by/2.0>, via Wikimedia Commons

 

When I wrote the recently article on the Michoacán Stream Salamander (Ambystoma rivulare), I remembered that readers of old books and papers, like Hans Gadow’s volume Amphibia and Reptiles in the Cambridge  Natural History Series first published in 1901, will be puzzled by the spelling of the generic name of mole salamanders, Amblystoma, rather than the one used today, Ambystoma.

In the last decades of the 19th century and the early part of 20th, Amblystoma was in general use for these salamanders of Canada, USA and Mexico, together with derivations like ‘amblystome’ as a common name. So why is the accepted name now Ambystoma?

It was Marcus Ward Lyon (1875-1942), medical man and mammalogist, who asked that question—with a certain degree of indignance— in a letter to Science in 1916:

Two letters were published in reply. The first, very long  and rambling, was from the etymologist Charles Payson Gurley Scott (1853-1936). He was pretty well convinced that Tschudi and/or the printer had made a slip of the pen (the lapsus calami) or a consistent typo since Amblystoma simply meant ‘blunt mouth’ and ‘Ambly’ was used in similar circumstances for other animals, the lizard Amblyrhyncus, for example. ‘Amby’ however had no meaning in the ancient languages. The second letter merely noted that another American author had used ‘Ambystoma’ as the correct form in 1909.

Leonhard Hess Stejneger (1851– 1943) had argued strongly that Tschudi's name should be used:

The scientific names of animals can be changed if there is evidence of a spelling mistake, however caused, in the original publication. One example is that of the American Alligator, originally named with the specific name ‘mississipiensis’ by Daudin in 1802 but later corrected to ‘mississippiensis’. However in the case of Ambystoma there was no evidence that Tschudi had suffered a lapsus calami and so the name used by Louis Agassiz (who must have also thought that Johann Jakob von Tschudi (1818–1889) had simply made an error) was replaced by the original, a case with so much Latin and Greek in play, of nunc pro tunc. Ambystoma was thus restored.

When Agassiz prepared his Nomenclator Zoologicus, a list of all names used in zoological genera  in 1842-1846 he was based in Europe. Both he and Tschudi were Swiss and I cannot help wondering if there was correspondence between the two along the lines of. ‘I say old chap I was just wondering if your name Ambystoma was the correct spelling for the name you intended?’ And if not, why not?

For my pennyworth, I would question why Tschudi would erect a genus meaning blunt-nosed for a  group  when European salamanders with which he would be familiar, Salamandra salamandra, for example, also have a blunt nose. Having seen Axolotls and Spotted Salamanders feeding on large earthworms I can see why stuffing the mouth might seem appropriate.

The world had other things on its mind in 1916 and well-known books, like Gadow’s were reprinted into the 1920s (with reprints appearing as late as 1968). Thus it is no surprise that, Amblystoma continued to be used well into the 20th century. Indeed I was surprised to see Amblystoma used by Malcolm Smith in his New Naturalist Series book, The British Amphibians & Reptiles, published in 1951. Perhaps those at the Natural Museum in London agreed with Agassiz and carried on regardless. Or perhaps that was a lapsus calami.

Lyon MW. 1916. Ambystoma not Amblystoma. Science. 43, 929-930.

Scott CPG. 1916. Amblystoma not Ambystoma. Science. 44, 309-311.

Stejneger L. 1907. Herpetology of Japan and Adjacent Territory. Bulletin of the United States National Museum 58, 24.

An Endangered Salamander in Mexico: The Toluca Axolotl or Michoacán Stream Salamander

As if a morning in Mexico gazing at a wildlife spectacular—one of the sites where Monarch butterflies gather in their millions from all over North America east of the Rockies to overwinter—was not enough, a short distance away from the entrance to the Monarch Reserve at Sierra Chincua and at an altitude at just over 10,500 feet (3220 metres) was another gem. A slowly-moving shallow stream across a cow field has a population of the endangered Michoacan Stream Salamander, and a number were out and about and feeding while we were there at the end of January.

We could see the salamanders moving about on the bottom of the slightly cloudy stream and had I not picked up the wrong-sized polarising filter for the camera lens when packing I would have better photographs and video to show. We were able to see them feeding on small invertebrates in the water and of blowing water from their lips to stir up the sand and reveal hidden prey.

Ambystoma rivulare, the Michoacán Stream Salamander or Toluca Axolotl

These salamanders are said to be like their close relation the Axolotl (Ambystoma mexicanum) in being neotenic but also capable of metamorphosis and breeding as transformed adults.

The evolution and taxonomy of salamanders from the Trans-Mexican Volcanic Belt is complex. It does not help when two species are sometimes known by the same common name, as in this case. However, it is clear from distribution maps and publications from Mexico on the fauna of the Monarch reserve that the species here is the Michoacán Stream Salamander, Ambystoma rivulare, which in Mexico is also being called the Mountain Stream Salamander. The distribution of A. rivulare in fact straddles the state border between Michoacán and Mexico (the latter a state within the country of Mexico).

Range of Ambystoma rivulare (from IUCN Red List) with the site we visited shown by the red dot

Ambystoma rivulare seems to have acquired another common name amongst Mexican scientists. The authors of a number of research paper call it the Toluca Axolotl, Toluca being the nearest city (and capital of the State of Mexico) to the sites where it can be found.

Confusion can arise because another species is also known as the Michoacán Stream Salamander. Thus two species with that same common name are listed in the IUCN Red List. The other species is Ambystoma ordinarium. However, that one is found further west than A. rivulare according to the distribution maps in the IUCN Red List.

The population of A. rivulare is reckoned to have decreased in recent years, particularly since non-native trout have been introduced to the streams for anglers.

None of the distances between the distributions of Ambystoma species in the Volcanic Belt is very great. Thus the ranges of the Axolotl (Ambystoma mexicanum), A. rivulare and A. ordinarium all occur within 100 miles, illustrating the adaptive radiation that has occurred in this volcanic region of central Mexico, producing what seems to be more than 10 species, each confined to a relatively small area.

The term ‘mountain stream’ suggests to me fast-moving water tumbling over rocks, quite unlike the habitat of A. rivulare that we were in, a stream across a flat field at high altitude. The only way to describe it was that it reminded me of the stream in which we used to catch Common Newts in the lowlands of the East Midlands 70 years ago.

Google Earth view of the site. The stream enters the pasture from the pine forest (top left)

Along with the Monarchs and a good lunch we had what can only be described as a grand day out—tempered only by the dreadful dinner later. 

The Tawny Mime - an uncommon butterfly in Hong Kong

AJP was walking over Tai Mo Shan in Hong Kong last month when he came across this butterfly. It is a Tawny Mime (Papilio agestor), a swallowtail. It is an unusual find in Hong Kong and the first specimen recorded there was found in 2002. The general description throughout its range in Asia is that it occurs in hilly regions - the location in Hong Kong fits accordingly. AJP also noticed that it was incredibly aggressive, seeing off any other butterfly it came across. Again that behaviour of flying a beat for long periods while attacking other large insects it encounters is characteristic of this species. 

The family connexion between ‘the father of American mammalogy’ and Hong Kong Country Parks

I have known the name Merriam for 65 years since I kept a Merriam’s Kangaroo Rat (Dipodomys merriami), a wonderful animal, for several years. For anybody unfamiliar with kangaroo rats, they are essentially bipedal, parallel evolution versions of the jerboas. Until I was looking up pocket gophers a few weeks ago after out trip to Mexico I had not given Clinton Hart Merriam much attention. To my surprise I found there was a family connection to an important figure in the conservation of Hong Kong and world wildlife in the latter half of the 20th entry.

CLINTON HART MERRIAM (1855-1942) is remembered as important figure in American mammalogy. However, he had wide interests in just about anything alive before while working as a family doctor. By what was clearly a bit of political help from his cousin, a senator, and his friend, Theodore Roosevelt, he became the first chief of what became Division of Biological Survey of the US Department of Agriculture, a post he held for 25 years. In later life though he abandoned these pursuits for ethnology, having become fascinated by the indigenous inhabitants of California and the western parts of the USA.

These days Merriam is often referred to as an out of control ‘splitter’ and erector of new species. Thus he described 86 species of jus the one Brown Bear in North America. Clearly though Merriam was important in both the natural history of North America and in the conservation of wildlife.

Clinton Hart Merriam

It was Merriam's grandson, LEE MERRIAM TALBOT (1930-2021) who was to have such an important role in world wildlife conservation, becoming head of IUCN from 1980 to 1983 and leading key pieces of U.S. and international legislation. Talbot was the action man of conservation in the early 1960s while working for IUCN, researching the status of endangered animals and habitats in many parts of reports while producing reports for governments and conservation bodies. In that work he worked closely with his wife, Marty (born 1932). South-east Asia was a particular focus in the early 1960s and his report for the Agriculture and Fisheries Department of the Hong Kong Government was produced as the result of, what I read, was a three-week trip in 1965.

Reports indicate that the work for the report was not without incident:

“While conducting an aerial survey for the government of British Hong Kong in the early 1960s, his plane experienced mechanical problems, and crash landed in a harbor, hitting rocks and pinwheeling through the cold, frothing water,” explained his son Russell. “He swam to safety, later describing in vivid detail the difficulty of determining which direction was up, while escaping the still-tumbling wreckage.”

Lee Merriam Talbot
from HERE

Lee Talbot was also a racing driver. He began at the age of 18 and had his last race at the age of 87. It seems 1965 was an unfortunate year. An account of motor racing accidents shows that in 4 July 1965 while he was based in Hong Kong he walked away from an accident that may well have killed him. In a Formule Libre race in Malaysia his Lotus Super-7 ran off the road and overturned into a drain, pinning Talbot under the car. His high quality helmet kept the weight of the car off his head. Another driver was killed in the same race

The Talbots’ work in south-east Asia all came together at a conference in Bangkok on 29 November-4 December 1965 organised by IUCN. The Talbots edited the proceedings which appeared in 1968. By the time the conference was held he had been recruited as ‘Smithsonian Field Representative for International Affairs in Ecology and Conservation’ in Washington DC.

Nothing happened to implement the Talbots’ report until the 1970s. The Hong Kong government was soon dealing with the communist-inspired riots of 1967 and their aftermath. The administrators of the time were not known for alacrity in their attitude to change and it was not until Murray MacLehose was appointed governor in1971 that much was done. Then work started on the Country Parks for recreation and conservation envisaged in 1965. The Country Parks Ordinance came into effect in 1976 with the first parks officially designated in 1977. At present there are 24 covering an area of 440 square kilometres.

Reading between the lines I suspect the Talbot report was the necessary icing on the cake of a movement within Hong Kong to create areas suitable for recreation and conservation. The pressure to obtain an external expert report seems likely to have come from individuals within the Hong Kong government itself, from the several amateur societies concerned with wildlife and the countryside (of which government officials were also members) and members of the university (there was only one then) staff from the botany, geography and zoology departments. Plans for some protected areas/parks had already been drawn up by the time the Talbots arrived for their short visit.

In 2013 Lee Talbot himself explained (HERE) the intergenerational interest in wildlife and conservation:

I grew up in a conservation family with endangered species conservation as part of my heritage. My maternal grandfather was Dr. C. Hart Merriam, founder and first head of the Biological Survey which became the U.S. Fish and Wildlife Service. A pioneer ecologist, he may be best known for the life zone theory of distribution of plants and animals which was a foundation of ecology for perhaps a half century. My mother was an ethnologist and naturalist, very concerned with conservation. And my father, M.W. Talbot, was a pioneer range and wildlife ecologist who, after years of field work in the southwest, was director of the California Forest and Range Experiment Station, the research branch of the U.S. Forest Service, and a professor at University of California, Berkeley. A lifelong conservationist, in 1924 he worked with his Forest Service friend, Aldo Leopold, initiating the Gila, the nation’s first wilderness area; helped with the early days of the Wilderness Society; was a founder of the Society for Range Management; and constantly worked for conservation and science-based sustainable management of the nation’s forests and rangelands.

     Hiking, camping and pack trips in the wilderness with the wonderful conservation and ecological insights and guidance of my parents was an integral part of my early years….

The Country Parks of Hong Kong in 2025
from HERE

Who connects a tropical lizard to a Hong Kong landmark, the U.S. president’s office…and a misspelt name?

For the answer let’s start in Washington D.C. where the president sits at a desk given to him by Queen Victoria. Her Majesty’s Ship Resolute was abandoned in the Arctic by the Royal Navy while searching for the lost expedition led by Sir John Franklin in 1854. The ship broke free from the ice the next summer and was found by an American whaler. It was repaired and returned to Britain by the U.S. government as a gesture of goodwill at a time when the president had been threatening British interests and wanting to annex Canada. In 1879 Resolute was decommissioned and three desks were made from its oak timbers. One of those desks was sent to Washington and was received at the White House on 23 November 1880.

The naval officer in charge of the five-ship squadron sent to search for Franklin was Edward Belcher.

Edward Belcher

It was Belcher, commanding HMS Sulphur, who was given the job of surveying Hong Kong harbour in 1841. He was present, having landed a day earlier, at the ceremony to take over Hong Kong on 26 January. Belcher’s Battery overlooking Hong Kong harbour was named after him when it was constructed in the 1880s. The battery was replaced by housing for government employees in the 1950s. That complex, Belcher’s Gardens*, in turn was replaced by six tower blocks with the name still retained—but oddly phrased—as The Belcher’s.

Hemidactylus brookii, a gecko, was thus named by John Edward Gray (1800-1875) of the British Museum in his catalogue of lizards of 1854. Two specimens from Borneo were given to the museum by Edward Belcher, presumably on his return to London on 31 December 1847. They were collected in 1843 when Belcher was in command of another survey ship HMS Samarang working on the north coast of Borneo. I will return to the gecko later. But first the man linking all these events and many more besides.

I have written previously on Belcher HERE. Edward Belcher was born in Halifax, Nova Scotia, in 1793. He joined the Royal Navy, at the usual age for an officer of 13, as a midshipman. His reputation can only be described as dichotomous. He was widely praised as a hydrographic surveyor, geographer and promoter of the natural sciences, his activities continuing after his retirement from the active navy list. By contrast, he was thoroughly loathed by almost all those he came into contact with whether his peers, his seniors or his juniors on board his ships.

Back with the gecko, here is Gray’s description of the beast.

In Borneo—and Sarawak in particular—Belcher was in close contact with and supporting the famous White Rajah, Sir James Brooke (1803-1868), in his efforts to clear pirates from the seas. Subsequent publications treat the gecko as having been named in honour of James Brooke. However it would appear that Gray did not know that Brooke had an ‘e’ on the end because he published the common  name as ‘Brook’s Hemidactylus’ and the specific name as ‘brookii’. In his 1854 catalogue he lists people who had donated specimens to the Museum. He shows Belcher in the list as well as ‘J. Brooks, Esq’. However, searching the volume there is no further mention of Brooks or Brooke. But we have ‘Brook’s’ and ‘brookii’ in the description of the gecko. Perhaps Belcher had intimated to Gray or his boss a patronym for Brooke woukld go down well but that Gray did not know of Brooke and how to spell his name.

Albert Günther in reviewing the reptiles and amphibians of Borneo in 1872 clearly realised the central role of James Brooke:

The first extensive collections received in Europe were from two localities :—1…. …2. From the principality of Sarawak, where Sir J. Brooke, Sir E. Belcher, and Mr. Low paid for a considerable period much attention to the fauna. The collections made by them were presented to the British Museum, and described partly by Dr. Gray, and partly more recently by the author in the Catalogues of the British Museum and in the ‘ Reptiles of British India.’…

Sir James Brooke
late 1850s. By Herbert Watkins
Natonal Portrait Gallery

Because of the evidence that the patronym was coined for James Brooke, some authors now use ‘Brooke’s Gecko’ for the common name.

But what of the gecko itself. When I was a lad Hemidactylus brookii was seen as a very widely distributed species of the tropics that, along with other species of gecko, had been transported by ship to other countries, like South America and parts of south-east Asia to which it was not native. Africa was regarded as its stronghold. In more recent decades H. brookii has been looked at more closely. Is it just one species or a number of different species that had either not been recognised or lumped into H. brookii by the early generations of taxonomists. And what a can of worms H. brookii has proved to be.

One of the problems stems from the way in which taxonomists worked. Taxonomist A would receive a specimen and decide if it was like anything else he had seen or been recorded. If not he would give it a new scientific name. Taxonomist B probably working in a different country would receive a similar specimen and, unaware of the work of A or disagreeing with him, would give it a different scientific name. Those coming later would gain access to both specimens or descriptions and declare them to be of the same species, or not. If the same they were synonymous and lumped into a single species, the name having priority being the earliest to have reached publication. The problem is that taxonomists usually gave no reason, either quantitative or qualitative, why they had reached their conclusion. Such decisions taken in the past, decided solely on morphological features of pickled specimens, have to be question in the light of the application of quantitative methods and/or non-morphological evidence like DNA. Depending on the ‘species concept’ being used by particular modern taxonomists, decisions are made on whether or not those old taxonomists were right or wrong, or perhaps I should say more likely to be right or more likely to be wrong since there is still disagreement as to what constitutes a species. Some of the concepts used seem nonsensical to outsiders like me and remind me of the drunk looking for his keys under a street lamp when he had dropped them further down the road in the dark on the grounds that it was easier to search where he could see. The complexity of what has been going on with regard to Brooke’s Gecko can be seen in the taxonomy section of the entry for the species in the IUCN’s Red List HERE together with the relevant references.

The present state appears to be that H. brookii has been split asunder. The African forms previously included in H. brookii have been assigned to a different but previously synonymised species (H. angulatus described by Edward Hallowell in 1854) leaving Asia as the natural range. Further studies on Asian specimens in museums indicate that other forms regarded as synonyms in the past should be reinstated while others should be brought back in.

Distribution of H. brookii from the IUCN Red List

Distribution of H. angulatus from the IUCN Red Data List

In a final twist to the story of Belcher, Brooke’s Gecko is now known to occur in Hong Kong. First reported in 1978, one location, St Louis School, is a few hundred metres from The Belcher’s; a large warehouse for imported goods once stood on the opposite side of the road.

Brooke's Gecko
from Hong Kong Amphibians and Reptiles - see below

James Brooke is buried in the churchyard of St Leonard's, Sheepstor, on the edge of Dartmoor: I tooke these photographs in September 2012:

Memorial Plaque to the Brooke Rajahs of Sarawak

*Belcher’s Battery and then Belcher’s Gardens were overlooked by the main teaching laboratory cum museum of the zoology department of the University of Hong Kong when it was in the now-demolished Northcote Science Building.

Gray JE. 1845. Catalogue of the specimens of lizards in the collection of the British Museum. London: Trustees of the British Museum/Edward Newman.

Karsen SJ, Lau M W-N, Bogadek A. 1998. Hong Kong Amphibians and Reptiles. Second Edition. Hong Kong: Provisional Urban Council. ISBN 962-7849-05-7