Paradox on the Queensland Frontier:
Platypus, lungfish and other vagaries
of nineteenth-century science

Part one

Libby Robin

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To facilitate downloading,
this paper has been divided
into parts one & two

I

Introduction
The central narrative of this paper is the 'discovery' by British scientist W H Caldwell that monotremes (platypus and echidna) lay eggs¹. The famous telegram 'monotremes oviparous, ovum meriblastic' (monotremes lay eggs of the same sort as reptiles), sent to the British Association for the Advancement of Science meeting in Canada in 1884, put to rest a debate about whether platypus laid eggs or had live young that had raged throughout the century. The story reveals much about the imperial shaping of scientific knowledge. Observers in Australia and Aboriginal informants, who had asserted that platypus laid eggs, had been disbelieved. 'Discovery' was reserved for Caldwell, a British scientist of impeccable scientific lineage.

This paper takes this well-known story and places it in a broader social context. What were the other scientific questions that drove Caldwell so far north in his collecting? And what was the significance of the cross-cultural frontier in his work? His 'discovery' depended upon astonishing support from Aboriginal assistants. The story suggests some new ideas about frontiers in the coalescing of scientific and cross-cultural frontiers in country that was already settled.

The Platypus Paradox
The platypus debate began in the eighteenth century. David Collins in 1797 saw 'an amphibious animal of the mole species… having, instead of the mouth of an animal, the upper and lower mandibles of a duck'.² George Shaw of the British Museum described the dried specimen he received in 1798 as 'of all the Mammalia yet known … the most extraordinary in its conformation; exhibiting the perfect resemblance of the beak of a Duck engrafted on the head of a quadruped'. The dried skin he examined is still marked by the scissors that he used to check that the beak had not been stitched on by a taxidermist.³

Shaw named it Platypus anatinus in 1799, and the German anatomist Blumenbach, independently named it Ornithorhynchus paradoxus in 1800.⁴ A genus of beetles already had been named 'platypus', so Blumenbach's Ornithorhynchus (genus name) was combined with Shaw's earlier species name, anatinus, to give today's scientific name Ornithorhynchus anatinus. Why was this animal not 'mallangong', (or another Aboriginal name)?⁵ The kangaroo (cunquroo) had defied description, so an Aboriginal name had been borrowed. The wallaby, koala and others had all similarly needed 'new' vernacular names. But the platypus, inexplicably, is known by a lost (Greek) scientific name that belonged first to a beetle. Until the mid-twentieth century 'duckbill' (a translation of Ornithorhynchus – literally, bird-nosed) was also popular.⁶ And the 'paradoxical' although lost to priority from the scientific literature stuck in popular consciousness. The scientific and the vernacular names have entangled stories.

Naming the platypus did not resolve where it belonged in the citadel of knowledge. Everard Home dissected a specimen in 1802, and was able to give a full internal description. Home noted its likeness to the echidna in having a common cloaca for reproduction and excretion: 'this tribe [has] a resemblance in some respects to birds, in others to the Amphibia'.⁷ Until 1824 evidence of mammary glands, the distinguishing feature of mammals, was undiscovered.⁸ Even when they were, the matter of whether the platypus and echidna gave birth to live young was unresolved, and this was seen to be critical to their place in the 'natural' world. It also became a question of nationalism. The French evolutionary thinker, Étienne Geoffroy Saint-Hilaire, ignoring the evidence of milk glands, separated the Order Monotremata, from 'true' mammals. He placed them halfway between mammals and reptiles. The British anatomist Richard Owen, disagreeing with the Frenchman's theories of evolution, saw them as definitely mammals, and therefore, he argued 'ovoviviparous' (the eggs were hatched inside the mother and the young born alive). Caldwell recorded three letters from colonial observers: John Jamison (1818), John Nicholson (c.1865) and George Rumby (1864), who claimed they had seen platypus eggs, but colonial evidence was not admissible to the faraway scientists of Europe, locked in bitter nationalistic rivalry.⁹

Whilst in Europe the platypus was dead and a puzzle, in the Australian colonies it was alive, familiar and personable. It was no longer perceived wet, camouflaged as 'a lump of dirty weeds', but increasingly its lovely fur was noticed. George Bennett in his Gatherings of a Naturalist in Australia 1860, described the thick fur as 'a beautiful adaptation to both the burrowing and aquatic habits of the animal'.¹⁰ He was reluctant to shoot the animal, trying to observe its behaviour alive.¹¹ The scientific and colonial encounter of the platypus was located in a particular place. The question of whose country yielded up the mystery of the platypus was not determined by the platypus or the people but by another 'missing link' in the evolutionary story.

Ceratodus (The Queensland Lungfish)
One of the last 'freaks' of Australian natural history to come to the attention of European science was Ceratodus (Serat-o-dus), the Queensland lungfish (Neoceratodus forsteri).It normally breathes through gills like other fish, but, when the oxygen levels in the water fall, it can rise to the surface and gulp air straight into its lung, an organ that other fish do not possess. The Australian lungfish is unlike lungfish in Africa and South America, in that it can live both underwater and on land. Fish with lungs were known only as fossils in the Northern Hemisphere at the time of the naming of Ceratodus, so the Queensland specimen was immediately dubbed a 'living fossil'.

Ceratodus had an immediate place in the history of ideas. Its relevance to debates about Darwinian evolutionary theory was obvious. Natural selection depended on continuities, but known classes of animals lacked 'missing links'. Fish, amphibians, reptiles, birds and mammals seemed discrete classes (apart from such absolute anomalies as the paradoxical platypus). Ceratodus, however, as a lung-breathing fish, was clearly intermediate between fish and amphibian. It was not unknown in 1870, only unknown to science. Not only Aboriginal people, but also the Mary River and Wide Bay district settlers ate it, calling it 'Burnett Salmon' for its pink flesh.

'Considering that the fish is not uncommon and has for some years been used as an article of food,' wrote Alex. M Thomson, Professor of Geology at Sydney University to Sir Richard Owen in 1870, 'it is surprising that it had not fallen into scientific hands much earlier'.¹² Owen, as Superintendent of the Natural History Collections at the British Museum, was in a position to dissect the fish and determine its place in the citadel of knowledge. Only at the heart of Empire were there sufficient type specimens of fish to decide where this one fitted. Gerard Krefft's independence in publishing in Australia – in the Sydney Morning Herald, of all places – suggested a growing bloody-minded independence in the hearts of colonial scientists at arms' length from good specimens.¹³ Ronald Strahan records that Gerard Krefft, the Director of the Australian Museum, had identified the fish as interesting when seeing so-called 'Burnett Salmon' being prepared for the table at the home of Mary River squatter and later New South Wales Minister for Lands, William Forster.¹⁴

Albert Günther undertook a full anatomical analysis of Ceratodus at the British Museum. Gerard Krefft's use of the name Ceratodusshowed that he was well aware of the fossil fish of the Northern Hemisphere and recognised that the 'new' fish had an ancient lineage. Even so Günther's anatomical description of the fish as an 'intermediate form' between fish and amphibians excited Krefft. In July 1870 (some time before Günther's paper on the anatomy of the fish was published), he wrote 'your Ceratodus forsteri if true a greatest discovery …[I am] amazed at it.'¹⁵ Although Krefft had given the fish its name, his use of the pronoun 'your' suggested that he was giving Günther credit for seeing additional significance in the specimen.

Why had Ceratodus been overlooked until 1870? There is no doubt that amateurs will preferentially collect warm-blooded species over cold, and obvious curiosities rather than technical ones. Perhaps the anti-evolutionary bias of the scientists contributed to the lack of interest. Mulvaney and Calaby commented of this period that: 'It was rather remarkable that the members of the Australian scientific establishment almost to a man…were vocal opponents of Darwin's ideas on the origin of species by means of natural selection.'¹⁶ Krefft himself ran foul of anti-evolutionary forces with his Trustees.¹⁷ The bitter battle that ensued after Krefft's dismissal put paid to further research papers from the Australian scientist most sympathetic to Darwinian evolution.¹⁸

I want to leave the hypothetical might-have-beens and return to the reasons for 'discovering' Ceratodus in 1870. The limitations of its habitat and the narrowness of its geographical distribution made its discovery by a scientifically-literate observer unlikely. The relevant part of Queensland could hardly be said to be new to European eyes at this time, but it was probably still fair to call it frontier country. Indeed the idea of a continuing frontier in Queensland dies hard. Wide Bay had been surveyed in 1848, and there had been 'settlement' up both rivers. Once 'exploration' finished, good scientific observers moved on to other pristine sites, leaving the settlement frontier to squatters and adventurers, whose natural history observations were untrained. When squatters at the frontier did make an observation, scientists were slow to believe them. William Forster had described Ceratodus but had been disbelieved by Gerard Krefft until 1870, a point Krefft confessed in his letter to the Sydney Morning Herald, where he attempted to make belated amends to the now important Forster as well as describing the new fish.

Gold and adventure seeking
The Gympie gold rushes of 1867 attracted a large population of adventurers. 'Gold upheaves everything, and its disruptions are like that of an earthquake', wrote Anthony Trollope travelling in Queensland in 1873.¹⁹ Trollope was as much concerned about the upheaval of morality as of the Queensland soil. The attractions of what we might call now 'boys' own adventures', were celebrated by the Englishman, Arthur Nicols in his fictional account Wild Life and Adventure in the Australian Bush: Four Years Personal Experience.²⁰ Writing eloquently of the 'noble territory of Queensland', Nicols boosts the frontier as a place for personal and financial development. 'These resources are waiting development at the hands of vigorous manhood which the upper and middle classes can contribute in abundance towards the making of this part of the Queen's realm'.²¹ His natural history observations are excellent and his displays a high level of curiosity about the platypus and echidna specimens he shot, including dissecting them, preserving the skins and eating them. Nicols' hero, Harold, tells a hunting story all about a man and his dog, Don and the hunt for an elusive platypus.²²

Because he worked within a particularly British sort of hunting ethic, Harold did not use Aboriginal collectors. That would be cheating.²³ The difficulty of catching the platypus made it a worthy 'trophy', despite the fact that it was not a lion or an elephant. The platypus for Nicols functions as a drawcard for young men of Empire with hunting aspirations. Australia's kangaroos and other marsupials did not generally carry the excitement of the wild animals of Africa. Nicols suggests that here was one that might arouse the sort of excitement where 'away flies conscience, philosophy and all such abstract considerations', in short, a manly challenge for the imperial hunter. ²⁴

Science and frontier life
The difficulty for scientists working in Queensland was, perhaps, to make it clear that their task was morally more worthy than that of frontier-adventurers. They would have been largely sympathetic with the evocation of empire, class and gender portrayed by Nicols. The difference for them was that imperialism was mediated by science, and science was in service to empire. British scientific visitors, in particular, were clearly fearless about taking very large numbers of specimens. But the scientists' virility was tied to hunting for knowledge rather than hunting for trophies. Such hunting demanded that they tap into the best local knowledge sources, including Aboriginal ones.

The young German embryologist, Richard Semon in his popular account of his travels in Queensland, announced the purity of the scientific purpose of his journey:

In June 1891, when I set out on my scientific journey, nothing whatever had been recorded with regard to the development of Ceratodus. Concerning oviparous mammals [there were] no developmental facts but that of their laying eggs, and an interesting observation about the teeth of young Ornithorhynchus…Thus it was that I chose Australia as my first and my main field of action, and within Australia those quarters which harboured the animals chiefly exciting my interest.²⁵

It is an account of 'an expedition' [not for mere adventure but] 'destined to bring some Phylogenetic Problems nearer their solution'. Before leaving Jena, Semon had precisely identified the Ceratodus territory where he would make his home in Australia:

[O]nly a brother naturalist will sympathise with me, when I own that an almost solemn feeling overcame me, on starting from the little station of Maryborough on the morning of 24^th August, I began my pilgrimage to land sacred to the zoologist.²⁶

W H Caldwell had made the scientific expedition internationally famous for this region, and Semon wasted no time in directing his attentions to the area where Caldwell's collecting had been most successful.

I now want to turn to the central story of this scientific frontier, the Ceratodus-driven collecting expeditions of Caldwell and his Aboriginal companions in the 1880s, which resulted in the famous telegram to the British Association about the platypus. It is very difficult to infer the Aboriginal perspectives from Caldwell's brief account alone. The fact that Semon went to precisely the same area only seven years later, however, and was introduced to Aboriginal collectors by the same squatter allows us to draw on his much fuller account of the Aboriginal people to supplement Caldwell's remarks.

Mr Caldwell's travels
William Hay Caldwell's Cambridge lineage was impeccable. The department he came from led the world in embryological research. He distanced himself from the 'early days of Darwinism' where 'it was hoped to get a pedigree for every animal'. 'Now that all biologists are Darwinists,' he declared to the Royal Society of New South Wales on 17 December 1884, where in fact there were very few Darwinian sympathisers, 'pedigree-hunting has gone out of fashion'.²⁷ Perhaps he was aware of the lack of sympathy to Darwinian biology in Australia and letting the colonials know they were out of step. He did not bother to write up this extempore talk, leaving it to someone else, in all probability a colonial not sympathetic to Darwinian biology, to report from his notes. Caldwell ultimately published so little that this second-hand account is crucial to gaining an idea of the state of his mind when in Australia.

Caldwell's work in morphology was to observe the minute differences between organic beings at various stages of development, in the belief that the patterns of evolution may be reflected in the patterns of individual development. His teacher at Cambridge, Professor Francis Maitland Balfour, had suggested in 1882 that Caldwell should consider travelling to Australia to work on the development of Ceratodus and 'the peculiar Australian mammalia'. Balfour, elected to a Fellowship of the Royal Society aged only 27, was taught by Professor Michael Forster. Forster in turn, was taught by T H Huxley, Darwin's most outspoken advocate. Caldwell benefited from the strong Darwinian lineage of Cambridge, but it was also a burden to him.²⁸ In 1882, Balfour, aged only 31, was killed in a mountaineering accident and a travelling studentship was endowed in his memory. Caldwell, Balfour's own student, working on a task assigned by Balfour, was the obvious first recipient of an 'instrument by which [Balfour's] memory was to reach beyond Cambridge and encompass the world for Darwinian biology'.²⁹ In addition to a personal salary from the Balfour Trust, Caldwell brought with him grants totalling £500 from the Royal Society (of London) towards the cost of equipment. Both Cambridge University and the Royal Society awaited the results of their investments.

George Bennett had studied the platypus mostly in New South Wales, and Caldwell, guided by this, determined to start in Sydney on his arrival in Australia in September 1883 and work over the platypus country inland. He had not counted on the skin-hunters. The trade in platypus skins had escalated dramatically in the two or three decades since Bennett's trips. By late in the nineteenth century, platypus rugs of forty and more pelts were being stitched together.³⁰ 'I wasted a fortnight trying to obtain information in Sydney as to where the animals were to be found in sufficient numbers for my purpose'.³¹ By mid-October, Caldwell had given up on platypus, and moved his attention to koala and wallaby, which were just beginning to breed. This material gave Caldwell new information on foetal membranes, and he sent home an account that was published in 1884 in the Quarterly Journal of Microscopal Sciences.

In April Caldwell went north to the Burnett River district to find Ceratodus, and noticed when he arrived that as well as Ceratodus with ripe spermatozoa, both echidna and platypus were numerous in the area. He decided to stay there for the monotreme breeding-season to try to get both Ceratodus and monotremes in the same year. 'The Burnett district', he wrote, 'presented the further advantage of possessing a considerable number of black natives. I afterwards found that without the services of these people I should have had little chance of success'.³²

Caldwell realised that to work with Aboriginal people he would have to create an independent camp. He set up out of Gayndah, away from the town, near the river where Ceratodus lived with provisions for his collectors. This was probably on the advice of district squatters such as W F McCord, who gave similar advice to Semon seven years later. McCord also recommended 'Frank' an Aboriginal from Gayndah to Semon as 'best adapted as to act as an agent between me and the blacks, to explain my wishes to them, and to be of help in my searches for the desired animals'.³³

It was probably no coincidence that the enterprising Frank turned up on Semon's coach from Biggendon to Gayndah, and became 'the first black who crossed my path'. Semon came to have reservations about Frank, however, and 'refused his services during my second stay in the Burnett.'³⁴ The majority of workers in the new science industry were Aboriginal, because of the particular skills required, but Caldwell reported that he employed some 'white navvies' to dig up platypus burrows because the Aboriginal team was reluctant.

Caldwell opened bidding with an offer of £10 to 'anyone who would show me Ceratodus spawn'. Once he had camp set up, he spent 'many hours in the water' in June and July hunting everywhere for the eggs of Ceratodus. Meanwhile:

the blacks began to collect Echidnaand very soon I had segmenting ova from the uterus. In the second week of August I had similar stages in Ornithorhynchus,but it was not until the third week that I got the laid eggs from the pouch of Echidna. In the following week (August 24) I shot an Ornithorhynchuswhose first egg had been laid; her second egg was in a partially dilated os uteri'.³⁵

This he described as 'a lucky chance'.³⁶ To kill a female platypus at the point betweenlaying her twin eggs gave him the crucial information about the stage of development at which the eggs were laid, a stage he described as 'equal to a 36-hour chick'.³⁷ The timing of the capture of this specimen was absolutely crucial. On 29 August he sent in the telegram 'Monotremes oviparous, ovum meriblastic' to a neighbouring station. The telegram was delivered to Professor Archibald Liversidge at the University of Sydney, who in turn sent it to the British Association at Montreal. Less than a week later, on 2 September 1884, Dr William Haacke from the South Australian museum was able to give evidence of egg-laying in monotremes, displaying an egg from an echidna's pouch at a meeting of the Royal Society of South Australia.³⁸ The Norwegian, Carl Lumholtz also claims to have heard reports of echidna's eggs, and was in pursuit of them at the time he had to leave Australia, being convinced that 'the reports I had received from the blacks corresponded with the facts'.³⁹ Caldwell's ability to tap the very consciously international audience in Canada was critical to his fame. The mystery was ripe for solution, but the drama of the telegram gave Caldwell an edge in the race.

Caldwell himself apparently hardly celebrated his historic moment. He was still anxious about that fish, and one can read in his words the pressure and burden that the Cambridge expectations had given him: 'Meanwhile I had never relaxed my efforts to find Ceratodus; but after four months I was beginning to despair of success'.⁴⁰ He finally found his Ceratodus eggs in September, and at this time employed some fifty 'black retainers'. Women were given the responsibility of trawling the river weed for Ceratodus, whilst men collected echidna, a favourite food. 'It was only occasionally, and then with great difficulty, that I persuaded them to dig for Ornithorhynchus. Not only the blacks, but their dogs, refused to eat the animal'⁴¹ It is yet another paradox of the platypus that the mystery of its egg-laying habits was solved with the assistance of Aboriginal collectors who themselves had no reason to hunt for the animal. This contrasts sharply with the earlier experience of George Bennett with Goulburn Aboriginal collectors. Bennett wrote:

The eyes of the aborigines, both young and old, glistened, and their mouths watered, when they saw the fine condition of the young Mallangongs. The exclamations of 'Cobbong fat' (large, or very fat), and 'Murry budgeree patta' (very good to eat), became so frequent and earnest, that I began to tremble for the safety of my destined favourites… But I was wrong in my calculation of the natives' power of resisting temptation, for they brought them all home safe...⁴²

Caldwell gained the critical knowledge and the necessary specimens from Aborigines for whom the platypus was not good eating. Neither was it a feature of their art, as it was for the Aborigines of the Kuring-gai Chase area, where a platypus is deeply engraved in a granite rock formation.⁴³ Burnett River Aboriginal people in large numbers assisted not only Caldwell but also Semon in obtaining the long embryological series critical to their science.

Continue with part two of this essay.

Author's biographical note and relevant links can be found at the conclusion ofpart twoof this essay.

Footnotes for Part One
¹ Elements of this argument will appear in a longer paper entitled 'The Platypus Frontier: Eggs, Aborigines and Empire in nineteenth-century Queensland' in Deborah Rose and Richard Davis (eds) Dislocating the frontier: essaying the mystique of the Australian outback.2001 (forthcoming). The longer paper focuses more closely on the nature of the frontiers. I am grateful to Deborah Rose and Richard Davis for comments on this paper.
² Charles Barrett, The Platypus, Melbourne, Robertson and Mullins, 1944, p. 12
³ Harriet Ritvo, The Platypus and the Mermaid and other Figments of the Classifying Imagination, Cambridge Mass, Harvard University Press, 1997, p. 4
⁴ George Shaw, The Naturalist's Miscellany: or Coloured Figures of Natural Objects Drawn and Described Immediately from Nature, Vol 10, June 1799 Fascicle, near plates 385 and 386. Johann Blumenbach, Abbildungen, Vol 5, Part 41, April 1800.
⁵ George Bennett noted that Mallangongand Tambreet are used in 'the Yas, Murrumbidgee, and Tumat countries' and 'Tohunbuck' at Goomburra, Warwick near Darling Downs in Gatherings of a Naturalist in Australasia,Milson's Point, Currawong Facsimile Edition, 1982 [Original edition, London, 1860] p. 97. Neither Richard Semon nor W H Caldwell record the Burnett River Aboriginal name for platypus.
⁶ The over-determined 'Duck-billed platypus' is still popular in Britain.
⁷ Cited in Ritvo, p. 7
⁸ J F Meckel had dissected mammary glands in 1824, but it was George Bennett's account of 'actual observation that milk is secreted from it' (letter to Owen, 4 February 1833) that was the basis for Owen's account of the glands to the Zoological Society of London in 1834. See Elizabeth Dalton Newland 'Dr George Bennett and Sir Richard Owen: A Case Study of Early Australian Science' in R.W. Home and S G Kohlstedt (eds.) International Science and National Scientific Identity, pp 55-74, esp. p. 68.
⁹ W H Caldwell 'The Embryology of Monotremata and Marsupialia – Part I', Philosophical Transactions of the Royal Society (B), vol. 178, 1887,463-85, esp. pp 467-8. He does not mention here that George Bennett also initially thought monotremes were oviparous. (See note 14)
¹⁰ Bennett, p. 97
¹¹ Australian Museum Archives: Series 37, Papers of George Bennett; Bennett's correspondence with Owen often refers to financial stringencies. (eg p. 54 – 'the Museum appointment has been made only £100 per annum & therefore anything but lucrative')
¹² Australian Museum Archives (AMA): Series 48, Gerard Krefft Correspondence 1861-78. Thomson to Owen 6 September 1870
¹³ Gerard Krefft, Sydney Morning Herald, 18 January 1870, p. 65, col. 5, Fig. 1-3; Krefft also published the description in the Proceedings of the Zoological Society of London, 1870, pp. 221-4, but the SMHwas first.
¹⁴ Ronald Strahan, Rare and Curious Specimens, Sydney, The Australian Museum, 1979, p. 29.
¹⁵ AMA 48: Krefft to Günther 13 July 1870. This is the only sentence in English in the letter.
¹⁶ D J Mulvaney and J H Calaby, 'So Much that is New': Baldwin Spencer 1860-1929, Melbourne, Melbourne University Press, 1984, p. 146.
¹⁷ Strahan, Rare and Curious Specimens, pp 33-34. George Bennett and W B Clarke resigned from the Museum over this.
¹⁸ G P Whitley and Martha Rutledge, 'Johann Ludwig (Louis) Gerard Krefft (1830-1881)' in Douglas Pike (ed) Australian Dictionary of Biography, Vol 5 K-Q, 1851-1890, Melbourne, Melbourne University Press, 1974, pp. 42-4.
¹⁹ Quoted in J W McCarty, 'Gold rushes', in Graeme Davison, John Hirst and Stuart Macintyre (eds.), The Oxford Companion to Australian History, Melbourne, Oxford University Press, 1998, p. 284.
²⁰ London, 1887.
²¹ Nicols, Wild Life and Adventure, p. vii.
²² Ibid, p. 347-8. The place of this platypus is probably south-west Queensland, not the Gympie area, but it is unclear, and because it is all 'fiction', conflation of places and events is possible. On the hunting ethic, see John MacKenzie, The Empire of Nature: Hunting, conservation and British imperialism, Manchester, Manchester University Press, 1988, esp. pp 25-53.
²³ The chief role of the one Aboriginal personality in Nicols' tale, Murray Jack, is to lead (and die in) a revenge battle against the Warrego people following the murder of a shepherd at an outstation. (see pp 315-39.)
²⁴ Harriet Ritvo, The Animal Estate: the English and other creatures in the Victorian Age, Cambridge, Mass: Harvard University Press, 1987, p. 267.
²⁵ Richard Semon, In the Australian Bush and on the coast of the Coral Sea. Being the Experiences and Observations of a Naturalist in Australia, New Guinea and the Moluccas. London: Macmillan & Co. 1899, p. 2.
²⁶ Semon, In the Australian Bush, p. 15
²⁷ W H Caldwell, 'On the development of the Monotremes and Ceratodus', Proceedings of the Royal Society of New South Wales, 18, 1884, pp 117-22.
²⁸ Roy MacLeod, 'Embryology and Empire: The Balfour Students and the Quest for Intermediate Forms in the Laboratory of the Pacific', in MacLeod and Philip Rehbock (eds), Darwin's Laboratory: Evolutionary Thought and Natural History in the Pacific, Honolulu, University of Hawaii Press, 1994, pp 140-165.
²⁹ Ibidp. 148. MacLeod records that £8,446 was subscribed to the scholarship.
³⁰ See Barrett, The Platypus,p. 17-19. The National Historic Collection of the National Museum of Australia has such a rug.
³¹ Caldwell, 'Embryology of Monotremata and Marsupialia', p. 464
³² Loc cit.
³³ Semon, In the Australian Bush, p. 17
³⁴ Ibid, p. 17
³⁵ Caldwell, 'Embryology of Monotremata and Marsupialia', p. 464
³⁶ Caldwell, 'On the Development of Monotremes and Ceratodus', p. 120;
³⁷ Caldwell, 'Embryology of Monotremata and Marsupialia', p. 464
³⁸ William Haacke, 'Exhibits', Transactions of the Royal Society of South Australia, Vol 7 (1883-4), p. 81
³⁹ Carl Lumholtz, Among Cannibals: An Account of four years travels in Australia and of Camp Life with the Aborigines of Queensland, London, John Murray, 1889, p. 329. Lumholtz was aware of Caldwell's work, so there may be some hindsight in this observation.
⁴⁰ Caldwell, 'Embryology of Monotremata and Marsupialia', p. 464
⁴¹ Ibidp. 465
⁴² Bennett, Gatherings of a Naturalist in Australasia,p. 132
⁴³ Barrett, The Platypus, rock art illustration.

Continue with part two of this essay.

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