John Samuel Budgett (1872–1904): In Pursuit of Polypterus

The patient persistence of his quest for the eggs of Polypterus under crushing difficulties forms one of the most courageous episodes in the history of zoology. —A. E. Shipley, “Biographical Sketch” (1907, p. 53). What drove scientists in the 19th and early 20th centuries to return again and again to the tropics, where the climate was appalling, risk of life-threatening illness a daily prospect, and the political situation fraught with danger? What possible questions could engender such stoic persistence? The goal of two turn-of-the-century biologists—John Samuel Budgett and Nathan Russell Harrington, both of whom died in the attempt—was to document the embryonic development of Polypterus, a fish found in the lakes, river margins, swamps, and floodplains of tropical central and western Africa and the Nile River system. Budgett, the more successful of the two, undertook four long, arduous expeditions between 1898 and 1903. The embryos he collected provided the basis for our understanding of Polypterus development for almost 100 years. This is the story of the pursuit of Polypterus and of the importance assigned to missing links between fishes and tetrapods. Discovered, described, and named in 1802 by Geoffroy St. Hilaire, Polypterus—a genus of 10 green-to-yellow-brown species, the largest of which can reach a meter in length (Figure 1)—posed a real conundrum for Victorian naturalists. What was one to make of an animal with a pair of lungs opening to the ventral wall of the pharynx, as seen in tetrapods and lungfishes and distinct from the single dorsal air bladder of teleost fishes pectoral and pelvic fins with fleshy basal lobes superficially resembling lungfish fins but with a fringe of numerous fin rays (lepidotrichia) not seen in lungfishes a subdivided, sail-like dorsal fin unlike that of any other fish a spiracle (persistent gill cleft), as seen in sharks and other cartilaginous fishes a thick armor of shiny scales covered, as are those of gars, with the specialized, mineralized tissue ganoine Was Polypterus then a bony fish, a cartilaginous fish, or a lungfish? Or was it perhaps even a primitive amphibian, in which case it would be a tetrapod and not a fish at all? In 1861 Thomas Henry Huxley created an order, Crossopterygii, to house those extinct and extant animals that possessed lungs and fleshy pectoral fins with lepidotrichia. Recognizing the specialized features of the two modern genera, Polypterus and Calamoichthys (Erpetichthys) calabaricus (the rope fish, which is longer and more slender than Polypterus), Huxley established a separate group (the tribe Polypterini) within the order Crossopterygii for them, noting Thus both ends of the Crossopterygian series appear, if I may use the expression, to be cut off from the modern representatives of the suborder; Polypterus being separated from those members of its suborder with which it has the closest zoological relations, by a prodigious gulf of time, and from the fossil allies which are nearest to it in time, by deficient zoological affinities. (Huxley 1861, pp. 445–446) 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 (Dean 1895, p. 149). Inevitably, the search for embryos of such species as Polypterus and the lungfishes led to far-flung regions of the globe, the homes of many of the species regarded as living fossils or missing links. Indeed, between 1877 and 1903 many embryologists set out from Europe for exotic locations in search of animal embryos. One of these was John Samuel Budgett (Figure 3). Nothing was known of the embryonic development of Polypterus, in large part because of the inaccessibility and inhospitability of the animal's habitat but also because recurring wars barred scientists from the interior of Egypt and the Sudan until the late 1890s. Collecting was dangerous, and it remained so for many years. This did not deter John Budgett, whose tenacity in what was to become “one of the most courageous episodes in the history of zoology” (Shipley 1907) brought success but cost him his life. John Samuel Budgett was born in Bristol on 6 June 1872. His father, W. H. Budgett, a keen microscopist, was a member of the Bristol Microscopical Society and served on the council of Bristol Museum. Budgett's was a childhood filled with the fascinations of the animal world. In the spacious grounds of Stoke House near Bristol, young John built aviaries, adapted outbuildings as homes for his pets, and organized a laboratory. He even had his own museum, complete with many animals he had meticulously dissected and stuffed birds displayed in their natural surroundings, not in a static museum setting. The museum was large enough to house his skeletal preparations, including a cow, a deer, and the family Shetland pony. Thus was laid the foundation for what was to become extraordinary skill in making anatomical preparations. It was said that his frequent visits to the Clifton Zoological Gardens to inquire about sick animals were prompted more by his desire to acquire specimens than by a concern for the animals' full recovery. He was skilled in keeping captive animals healthy, however. He was also a gifted draftsman and a fine watercolorist. Budgett began his formal study of zoology at University College, Bristol, having been much influenced by his father's many scientific friends, especially the then-elderly but still eminent anatomist W. Kitchen Parker, who for almost a quarter century spent part of each summer at Stoke House. So inspired was Budgett by Parker's work on the development of the skull that he began working on the topic himself, using his skills in dissection and specimen preparation. He even designed a device for aligning serial sections that enabled him to construct models of the developing skull. Budgett drew consecutive sections on plates of ground glass of known thickness, arranged the plates in series, then immersed them in a bath of oil, which rendered the glass translucent and allowed the outlines of the specimen to be seen as a solid figure (Figure 4). Budgett tackled all the major problems associated with generating three-dimensional reconstructions from histological sections, including how to line up the sections, a problem that bedevils us still. The young man, described as unassuming, modest, diffident, and charming—though subject to occasional bouts of severe depression—entered Trinity College, Cambridge, in 1894. With his prodigious knowledge of natural history and strong testimonials, Budgett was soon welcomed into the inner sanctum of Cambridge zoology. He was elected to the Cambridge University Natural Sciences Club in his first year, a rare honor indeed, for this exclusive club was limited to 12 undergraduate and a few graduate members. In August 1896, after completing part I of the Natural Sciences Tripos, Budgett embarked for Paraguay to collect lungfish embryos with John Graham Kerr (1869–1957), who had just completed part II of the Tripos. Kerr, a superb 19th-century zoologist who worked for six decades into the 20th century, had expressed his naturalist tendencies early, interrupting his medical studies at Edinburgh University to join the 1890 Page expedition to South America. (Kerr later published A Naturalist in the Gran Chaco [1950], a spellbinding account of this and subsequent expeditions and an adventure tale that ranks with Darwin's Journal of Researches for sheer excitement and impact.) The expedition was mounted by the Argentinean Navy to survey the Pilcomayo River from the Paraná River north some 500 miles to the Bolivian frontier. Because of the appalling conditions under which they had to be transported, many of the new species collected during the 3-year expedition were lost. Kerr did not resume his medical studies after he returned from South America in 1892; instead, he entered Christ's College, Cambridge, to read natural history. His studies on Nautilus, carried out as an undergraduate—the first “modern” account of the anatomy of this species—are important because they demonstrated the relationship between cephalopods and the rest of the molluscs. Such studies convinced Kerr of the benefits to be gained from studying the more archaic members of a group. It had been known since 1836 that the lungfish Lepidosiren lived in the Amazon, so when Kerr learned during his 1890 trip to South America that a soldier had caught an eel-like fish in the mud of the Pilcomayo River, which forms the border between Argentina and Paraguay, he wondered whether it might be a lungfish. Together, Bohl's [the German collector who found Lepidosiren in the River Paraguay in the Gran Chaco in 1894] discovery, my memory of the soldier's tale, and the fact that I had unique experience of life in the Chaco, decided me to give priority to the Lepidosiren part of my programme and to proceed the moment I had got quit of my Tripos to organize an expedition to the Gran Chaco [a vast area of plains, rivers, and swamps in central South America, which extends to Paraguay, Bolivia, and Argentina] for the purpose of tackling the Lepidosiren problem. (Kerr 1950, p. 172) As Kerr recounts, John Budgett was an unlikely companion on such an expedition: While engaged in planning my expedition there came to me one day a friend who said “John Budgett of Trinity is just dying to go with you.” I pooh-poohed the idea at first, for Budgett had no experience of roughing it in wild country, but his friends were persistent so at last I invited him to dine with me at the Café Royal and talk matters over. That talk, helped by an admirably cooked roast wild duck washed down by Clicquot–vin rosé, resulted in the settling of all my doubts, and the recruitment of a courageous, tough, and loyal comrade. (Kerr 1950, p. 172) After tea went out with gun across the swamp and had a fearful time, rushes had grown high, some had fallen horizontally, so that at each step one's feet were sawn across with fine teeth running along the under-surface of the midrib; socks were soon gone to shreds, and skin began to go. After half an hour's walk of this sort I reached the bird island, but was welcomed by such an attack from the garrison stationed there that I immediately turned and fled into the swamp again, fighting my way back to the Toldo [hut], until I arrived pretty well exhausted, having only shot one small bird. (Shipley 1907, pp. 10–11) It was a remarkable feat. To go straight as an arrow to the place where this almost unknown fish lives, to arrive at about the time of the breeding season, quite unknown before, and to collect and preserve all the delicate and varying stages of development within some seven the expedition at the first of Zoological there is which it even more and that is the of the when it The in by all of by and by and been but it was The success in this was to his skill in and his knowledge of the use of (Shipley 1907, pp. having been a in Budgett was to a in part II of the Natural Sciences Tripos in by his South he was by a desire to and embryos of Polypterus and to to The to of the of the to Budgett, Nathan an and a at University who was working on the and of had set out to Polypterus embryos. and a in at reached on 1898 in the of the the they spent in and and and and for on the most locations for Polypterus, there is no that Budgett and did Budgett the of the the of at a time, and a search that until 10 and them miles up the the to with at The two returned to the in with specimens of Polypterus, some of which they had to the fishes and other and a large of In late a on the and breeding of Polypterus to the which was in the completed zoological of The specimens allowed him to the to the lungs and the ventral opening of the lungs to the He that Polypterus was as well as any to be a lungfish. The and a to those of described an in the and provided on the Because the of all Nile fishes he that the breeding of Polypterus also the of the Nile that Polypterus an between fishes and and so could light on the of the In the of and in primitive skeletal it the fishes that were of the but from the conditions in Polypterus they were also in the of to to both and p. While was to Budgett, with and from the Zoological for his 1898 from for The With him on the he carried of Polypterus to the he had from of the Cambridge The of a of than 10 on side of the River, which This established after a with the in August and on each side by some of whom to the the to The was regarded as having the climate on the The was with of in to The was from most of the had died in and was no but was by a of which Budgett later by and the River, which is to Budgett's first expedition a of the time was spent at the on the River from which to The first Polypterus as the brought to Budgett, on was two on the of the and died (Shipley 1907, p. It almost to Polypterus in the Budgett found that the to the of by the the way to young June he had dissected many including with eggs in their His first at almost and Budgett swamps and a single developing he was not his his from occasional a to (Shipley Indeed, Budgett's few of the arduous conditions under which he even he was to with by the of to return to the Nile to Polypterus Harrington, and from had to to a small and on the of the the River, which just the some of In Budgett with and after only one in the the came down with what was described as Nile which and within his life. In Budgett arrived back in with a pair of Polypterus, which he to the Zoological Society on the two Polypterus and for at and even what he to be He described how the pectoral fins are in and how the air bladder as a to for up to a day in a He the breeding as having to swamps in June and in the of August and and return to the in and Budgett had returned to on the of the South he was keen to for in recurring of had his and he had to with the of the Cambridge University He the first of that in Budgett a trip to The in in June so as to arrive for what he to be the breeding season, which with the The season, which so is a for and that of were That when Budgett arrived at he found the as as a and in a and their including were their cut and their The and Budgett spent for He eggs and and A single from on this the basis of a to the zoology of the for the of on The specimen possessed a dorsal an skull and a single pair of with from immediately the spiracle (Figure Budgett the features of Polypterus but also to the sharks and rays in the pectoral and to the teleost fishes in the and to the in the bony He that were a central group of fishes but not to the Harrington, on the other hand, had them to lungfish and In Budgett was named of the Cambridge zoology museum, where he spent to a day anatomical specimens for the Budgett's to the after the of in the in with from the and a from the Zoological enabled him to return to Africa in what would to be trip in pursuit of embryos. The John a in who was the in Cambridge, Budgett to for Polypterus in which of Budgett in June so that his would with the This trip was spent in in and near the of the His of some four or two and four as well as a which Budgett to of the to time to the exotic and Budgett from to the Nile in but was he found many Polypterus, at again In his own in the I my attack on the Polypterus problem. I had species of Polypterus to work was after I up my and returned to having got a of on the life and of Polypterus, having seen of the and of the most river in the but having again in my to the early stages in the development of p. The early part of 1903 was spent working up his of fishes in the Cambridge of Budgett two to the Zoological Society on One described the trip to and the other was on the spiracle of Polypterus, which he was to and air from the lungs that a had brought back a young Polypterus from in the River Budgett to to on the he of in an to to on a The one of the in the from his own that it would be to embryos in than in The or the Nile Budgett set off in August he six in all with large August he had his most on I to the he the eggs of has seen them at the and of the of at of eggs to under in He they are in two or I until to go him for In the had small Polypterus brought with (Shipley 1907, p. August Budgett the the to be but had of the eggs were or in the brought him two more one of which had many in its of these were Budgett to that was which it is on Budgett eggs and and for the first time the embryonic development of Polypterus later he a to Graham Kerr p. that as it was then was being complete the and the into of almost The of the into the during of the and development of the were also the importance of his The been for some time the most important group the of the and the gained by Budgett in the working out of his in the be to with the greatest interest by all p. with a of eggs Budgett an series from the stages to early which he in or The to on his that for after his return to A to a friend on up both his and the I brought off the of about a and that which I been to for the four years. The interest to me of the day by and by I can as I my as to this extraordinary relations to other It almost is and are with tropical and by The for my in that I eggs is I be to my the of this is almost especially when to (Shipley 1907, p. The ends on the last day of His by of his not from the to that he had not been named of the Zoological Budgett returned to that I would not part with for a (Shipley 1907, p. After a few with in and with his in Budgett returned to Cambridge to work on his The first of which of those it on the that he his of the features of the embryos. the Budgett until his family but his He died on the day he was to his to the Zoological Budgett some eggs and embryos and he no not even a single It was to Kerr to the embryos and of there were in the series and the of were Kerr that as a has been found to be to give a pretty complete of the of development of a Crossopterygian and this has of been a to it is to of the that has been for (Kerr p. In a to the of the for the of in in and in subsequent E. all the that Polypterus is not a it within the the most primitive of the fishes This Polypterus from the line to that has no with Polypterus is a and a primitive the in his of early fish The of is a and even many a to Polypterus as a and it is so described in many a of fish the of this for many not a but an and can us about anatomy and pp. What of more recent studies of Polypterus and both the specimens collected by Budgett in their studies of the development of the skull and line system. are of some features of specimens collected by on the expedition of also in species of Polypterus been in there been only two subsequent of series, those by and a complete of all embryonic and stages by and and series enabled and to a major by Kerr with their study of the of The for series has from that which The embryonic and development of the the group of all other living is The of and stages may understanding of in the of these primitive and may as a for with the of and as well as to that of some of p. The study by and the early embryonic stages by Budgett and described by Kerr, was to be Polypterus development is much more than lungfish between the of Polypterus and the of that of other of were (Figure p. In most other the questions that prompted Budgett to go to such in pursuit of Polypterus embryos I my for from the Natural Sciences and of and the of University is Polypterus, with its fleshy fin lobes with many fin and sail-like dorsal fin (Dean of Polypterus Lepidosiren and a remarkable (Kerr John Samuel from Kerr Budgett's of his to the skull of a of are in A of Polypterus, by Budgett in the in the

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