[2] However, the long periods spent in swampy, mosquito-ridden places left him debilitated with malaria and he died of blackwater fever shortly after his return to England.

[2] Parker had studied the development of the skull and John began working on the topic himself, using his skills in dissection and specimen preparation.

The museum contained minute dissections, skeletons (including a cow and a deer) and stuffed animals all prepared by him and presented against natural-looking backgrounds.

[4] In August 1896, in his third and final year as an undergraduate, Budgett accompanied a newly graduated Kerr on an expedition to the Gran Chaco of South America.

He had conducted original research on Nautilus as an undergraduate and found that it provided a link between cephalopods and the rest of the molluscs.

Nevertheless, by the summer of 1897 Kerr and Budgett brought back to Cambridge a large supply of adult lungfish, with major embryonic and larval phases, so well preserved that the finest detail could be seen.

The difficulty in improvised rooms, worried by all sorts of insects, by torrential rains and occasionally floods, by inquisitive and highly suspicious natives...must have been enormous, but it was overcome.

[2] Kerr had designed an apparatus to reconstruct solid figures from a series of microscopic drawings which have been obtained from sections or slices of the original biological structure.

Budgett applied his ingenuity to make it easier to get the glasses in exactly the right position and his knowledge of reagents to choose an oil that allowed the maximum possible amount of light to pass.

[1] In 1861 Thomas Henry Huxley had created an order: Crossopterygii to house those extinct and extant animals that possessed lungs and fleshy pectoral fins with lepidotrichia.

[1] In 1895, Bashford Dean, a leading authority, wrote: "From their isolated position, these recent forms [Polypterus and Calamoichthys] become of extreme interest to the morphologist, and from the side of their development, when this comes to be studied, they are expected to throw the greatest light on the relations of the primitive Teleostome to the sharks and Dipnoans, on the one hand, and to the Ganoids on the other".

Although he had failed in his main goal he had gained experience; he had established the breeding period of Polypterus and he had collected information about the fauna of the Gambia.

[2] Quite independently, Nathan Harrington (1870–1899), an experienced field biologist and a doctoral student at Columbia University had already set out to find Polypterus embryos.

Harrington and Reid Hunt, a tutor in physiology at Columbia, reached Cairo on 26 May 1898 in the midst of the Anglo-Egyptian campaign against the Mahdi.

The two men returned to the United States in December with preserved specimens of adult Polypterus, fishes and other vertebrates, and a large collection of invertebrates.

In late December, Harrington presented a paper on the respiration and breeding habits of Polypterus to the American Morphological Society.

Harrington thought that Polypterus occupied an intermediate position between fishes and amphibians and so could shed light on the origins of tetrapods.

[1] In 1899 Harrington got permission to travel with Hunt and F. B. Sumner (also from Columbia University) to a small village and military post on the northernmost tributary of the Nile, the Atbara River, about 650 km south of Khartoum.

Instead he joined the Mounted Infantry section of the Cambridge University Volunteers, giving all his spare time to the training of men for the War.

[2] Budgett traced the development of Gymnarchus niloticus and made observations on other fish, frogs, mosquitoes and other insects.

[2] On 29 August he gave up and proceeded further north by boat, passing Wadelai, Nimule, Kiri, Legu and reaching Gondokoro on 22 September.

He did hope to apply for the job of Resident Superintendent so he and Shipley took a continental tour at Easter, 1903 visiting zoos at Paris, Frankfurt, Leipzig, Berlin, Hanover, Amsterdam and Antwerp.

He also says we owe to Budgett the first accurate knowledge of the urinogenital system of Polypterus and the demonstration that the crossopterygian fin is really a uniserial archipterygium as well as observations of the life-history and breeding habits of many tropical frogs and fishes.

[4] This contains all his papers as well as a biographical sketch by A. E. Shipley and assessments of his work by several authors In a report to the 1907 annual meeting of the British Association for the Advancement of Science, Edwin Stephen Goodrich amassed all the evidence that Polypterus is not a crossopterygian, placing it within the palaeoniscids, the most primitive of the ray-finned (actinopterygian) fishes.

[1][8] A 1997 study[9] confirmed the early embryonic stages drawn by Budgett and described by Kerr, although fertilisation was shown to be external.

Budgett J. S. (1899a) Notes on the Batrachians of the Paraguayan Chaco, with observations upon their breeding habits and development, especially with regard to Phyllomedusa hypochondrialis, Cope.

Budgett J. S. (1901b) On the breeding habits of some West-African fishes, with an account of the external features in the development of Protopterus annectens, and a description of the larva of Polypterus lapradei.

Bartsch P., Gemballa S., Piotrowski T. (1997) The embryonic and larval development of Polypterus senegalus (Cuvier 1829): Its staging with reference to external and skeletal features, behaviour and locomotory habits.

The Work of John Samuel Budgett, Balfour Student of the University of Cambridge: Being a Collection of His Zoological Papers, together with a Biographical Sketch by A .

The complete mitochondrial DNA sequence of the bichir (Polypterusor natipinnis), a basal ray-finned fish: Ancient establishment of the consensus vertebrate gene order.