938:. At last he began to get specimens with ova. He started artificial insemination and was able to watch the key formative stages, lasting about five days. On 9 September he wrote to a friend that he had fertilised about a thousand eggs and watched them develop in the way that he had expected. Sadly after a week, a fungus had attacked and they had all died before reaching full maturity. However he continued his work right through September and finally managed to preserve specimens of each stage of development including a few larvae.
419:
expedition at the first rank of
Zoological exploration. But there is another factor which makes it even more remarkable, and that is the wonderful condition of the material when it arrived.... 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. The success in this respect was certainly partly due to his skill in manipulation and his peculiar knowledge of the use of reagents.".
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to a piece of ground glass whose thickness bears a definite relation to the thickness of the section. Then the glass plates are placed in a bath of oil which eliminates the opacity of the ground glass and should provide a 3D image of the original 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.
33:
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but in vain...Budgett succeeded but for this success he paid with his life....from each of these rivers he brought home not only most valuable notes on the habits of the fishes he came across, habits which were then totally unknown...embryological material.... very important collections of the fishes
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but not developing ova. He tried artificial fertilisation and he tried holding fish in cages to breed but these methods failed. He was constantly battling against the heat and against illness. Towards the end of July he was confined to bed and on 28 July he started his return journey. Although he had
434:
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. This might for example be the whole or part of an organ of an animal. The principle is that each section is drawn on
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says, "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 months, placed the
411:
Conditions were difficult. At times the
Pilcomayo is 300 m wide, at times it is a trickle or stagnant swamp, marsh, lagoons, and dense tropical jungle. It has sand banks, confusing channels, and barriers of overturned and floating debris. Nevertheless, by the summer of 1897 Kerr and Budgett brought
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reviewed his work and wrote,"The breeding habits of fishes living between the
Tropics are among the secrets of Nature which are most difficult to unravel, and which have most taxed the acumen and patience of zoologists...Several attempts have been made with the object of procuring the development
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John was fascinated by the animal world, building aviaries and adapting outbuildings for a museum and a laboratory. 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. He visited
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Kerr, J.G., ed. (1907b). 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 . E. Shipley, F.R.S., and Contributions by Richard Assheton, Edward J.Bles, Edward T. Browne, J. Herbert
543:
In 1895, Bashford Dean, a leading authority, wrote: "From their isolated position, these recent forms 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
759:. It set out on 11 July, according to Budgett’s diary, "with forty porters, four askaris, two headmen and four boys." Budgett himself used a cycle where possible to give himself additional time to make observations. They reached the lake on 30 July and almost immediately found specimens of
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Parker had studied the development of the skull and John began working on the topic himself, using his skills in dissection and specimen preparation. He designed a device for aligning serial sections that enabled him to construct models of the developing skull. He became an expert in this.
431:(the Cambridge equivalent of Finals). He was not good at examinations but managed to get a Second. He then set to work on his South American frogs and completed his paper on them. He was a gifted draughtsman and watercolourist. The figures in his papers were both artistic and accurate.
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in 1894. With his knowledge of natural history and strong testimonials, Budgett was elected to the
Cambridge University Natural Sciences Club in his first year, a rare honour, for this club was limited to 12 undergraduates and a few graduate members. He was also elected a member of the
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A 1997 study confirmed the early embryonic stages drawn by
Budgett and described by Kerr, although fertilisation was shown to be external. Development was found to be much more rapid than for lungfish. They found close similarities between the ontogenies of
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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. Budgett’s skill in manipulation and his knowledge of reagents played an important part in this success.
842:. The committee was impressed by his experience of keeping live animals in captivity but chose an older man. 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,
173:). This is found in the lakes, river margins, swamps, and floodplains of tropical central and western Africa and the Nile River system. Zoologists at the time wondered whether it was a bony fish, a cartilaginous fish, a lungfish or a primitive
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Noack K, Zardoya R, Meyer A. (1996). The complete mitochondrial DNA sequence of the bichir (Polypterusor natipinnis), a basal ray-finned fish: Ancient establishment of the consensus vertebrate gene order. Genetics 144:1165–1180, cited by Hall
270:. For twenty-three consecutive years Parker spent a fortnight at Stoke Park. He was noted for his skill in preserving delicate skeletons, such as tadpoles and small birds, and he appears to have passed some of this skill to John.
966:. He held his own for a while. The blackwater fever receded but then he had an attack of malaria as well. He died on 19 January. This was the very day on which he had expected to report his work to the Zoological Society.
709:
In May 1901 Budgett was appointed
Assistant Curator of the University Zoological Museum. He made a number of beautiful preparations to demonstrate anatomy to students. He published four papers based on his expeditions.
628:. Their search lasted until 10 September and took them 377 miles up the Nile but they failed to find adults with mature eggs. The two men returned to the United States in December with preserved specimens of adult
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Budgett returned to the Gambia in the rainy season of 1900, arriving at McCarthy Island on 6 June. The country was uncharacteristically dry and politically in a disturbed state. On 19 June, news came that the
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Shipley says that
Budgett did experiments with reagents, materials and equipment which advanced zoological technique in the laboratory. He also says we owe to Budgett the first accurate knowledge of the
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often to obtain dead animals. He was skilled in keeping animals healthy, a good draftsman and watercolourist. In the summer he used to get up at 3 a.m. and walk miles to study animals and birds at dawn.
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was remote and inhospitable and partly because of recurring warfare until the late 1890s. Budgett worked to fill this gap for five years and eventually succeeded, but only at the cost of his life.
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acquired in the
American and African swamps. 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.
343:. He had concluded that there was a general case to be made for studying the archaic species in a group to shed light on the group’s evolution. It had been known since 1836 that the lungfish
266:, a successful wholesale grocery firm. Often he had professional scientists to stay during the summer and John was able to get informal tuition from them. Prominent among these was Professor
736:, this was the top studentship at Cambridge for a research student in Zoology. With its income, together with various grants, the trip became possible and Budgett set out in May, reaching
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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
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As a memorial, Budgett’s friend Graham Kerr edited a book. This contains all his papers as well as a biographical sketch by A. E. Shipley and assessments of his work by several authors
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and spending the winter in Cambridge. Roscoe was well informed about the local people and about the natural history. He convinced Budgett that the best place to look for developing
587:. He sailed on 19 October 1898, arriving on 3 November. His stay lasted eight months. He kept a diary and recorded many observations. While searching for the breeding place of
651:. Sadly in July, after spending only one night in the desert, Harrington came down with what was described as Nile fever. It rapidly worsened and took his life within days.
184:, zoologists were seeking to map the history of species and this primitive animal was a key part of the map. To find its place there, it was necessary to observe juvenile
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Budgett J. S. (1899b) General account of an expedition to the Gambia Colony and Protectorate in 1898–99. Proceedings of the Zoological Society of London 1899: 931–937.
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was a job which he had hoped for and it fell vacant but someone else got it. Over the Christmas holidays he spent time with Shipley in Cambridge and with his mother in
290:. However, he had to leave the College at about fourteen owing to severe headaches caused by an accident. He continued his studies with private tutors before entering
1203:(1908). On the systematic position of Polypterus. Report of the 77th Meeting of the British Association for the Advancement of Science (1907): 545–546, cited by Hall
632:, fishes and other vertebrates, and a large collection of invertebrates. In late December, Harrington presented a paper on the respiration and breeding habits of
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is a good actinopterygian and a primitive one". He also said, "The weight of Huxley’s (1861) opinion is a heavy one and even today many a text continues to cite
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Budgett’s ambition was to attack one of the great unsolved problems of zoology. Having watched Kerr work on lungfish in South America he resolved to work on two
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Budgett J. S. (1899a) Notes on the Batrachians of the Paraguayan Chaco, with observations upon their breeding habits and development, especially with regard to
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were large but impossible to catch. So he carried on to Lokoja. There he inspected the possible fishing places but was not impressed so he continued towards
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1895. Fishes, Living and Fossil. An Outline of their Forms and Probable Relationships p. 149. Delhi (India): Narendra Publishing House., cited by Hall
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on 16 June 1872. His parents were William Henry Budgett (1827–1900) and Ann (née Lidgett, 1839–1936). When John was two they moved to Stoke House and Park,
1188:(Cuvier 1829): Its staging with reference to external and skeletal features, behaviour and locomotory habits. Acta Zoologica 78: 309–328, cited by Hall.
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1039:
663:. He had always been patriotic and his travels had made him more so. He felt he should go to the front but he was liable to recurring attacks of
203:. He didn’t even have time to write a report. However he did leave a full set of drawings and specimens. It was left to his friend and colleague
763:. The females had mostly shed their eggs so he decided to proceed north where the season was later. On 22 August he stopped for several days at
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is not a crossopterygian, placing it within the palaeoniscids, the most primitive of the ray-finned (actinopterygian) fishes. This removed
1151:
Budgett J. S. (1901b) On the breeding habits of some West-African fishes, with an account of the external features in the development of
1036:
931:. On 5 August he moved on to Dakmon, arriving two days later. This too was unpromising, so he set back to Assé, arriving on 12 August.
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and of urodele amphibians and other 'primitive' amphibians. Other questions that Budgett was hoping to answer are still unanswered.
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In August 1896, in his third and final year as an undergraduate, Budgett accompanied a newly graduated Kerr on an expedition to the
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females (locally called "Intontos") laying or having laid eggs. He was certain that the fry must swarm in the floating vegetation (
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1094:... is not a crossopterygian, but an actinopterygian and hence can tell us nothing about crossopterygian anatomy and embryology."
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958:, returning to Cambridge early in January. By 9 January he had finished his drawings of the external features of the developing
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shortly after his return to England. This happened on the very day that he was supposed to deliver a lecture on his work to the
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1233:(1982). Morphology and interrelationships of primitive actinopterygian fishes. American Zoologist 22: 241–260, cited by Hall.
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in central South America, Budgett did useful work on the amphibians of that area and discovered two species of a new genus
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It was very wet but he worked hard, fishing and also collecting and interviewing local fishermen. His guide knew about
262:. He was a keen amateur student of zoology and encouraged his children to study it. Professionally he was a partner in
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delta and Budgett thought that if he went there he might finally be able to observe the development of these larvae.
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occupied an intermediate position between fishes and amphibians and so could shed light on the origins of tetrapods.
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so when Kerr learned in 1890 that a soldier had caught an eel-like fish he wondered whether it might be a lungfish.
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and made observations on other fish, frogs, mosquitoes and other insects. He returned in August. He had caught 127
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On his return to England, Budgett was suffering bouts of malaria and his hands had sores from constantly handling
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He decided to trade on the fact that his Balfour Studentship was named after the brother of the Prime Minister,
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1223:
Kerr, J.G. (1950). A Naturalist in the Gran Chaco. Cambridge (UK): Cambridge University Press, cited by Hall.
950:. However he returned to Cambridge and began to work out his material. The post of Resident Superintendent at
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Quite independently, Nathan Harrington (1870–1899), an experienced field biologist and a doctoral student at
659:
When Budgett returned to England in the Autumn of 1899, he found the country in a turmoil on the eve of the
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Budgett J. S. (1903a) Account of journey to Uganda. Proceedings of the Zoological Society of London: 2–10.
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1127:, Cope. Also a description of a new genus. Quarterly Journal of Microscopical Sciences 2 : 305–333.
294:. There he received help in the formal study of zoology but much of his practical skill was self-taught.
291:
884:. The answer was negative but Balfour very generously paid for Budgett’s passage out of his own pocket.
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were caught but they had already shed their eggs. He returned to Khartoum on 10 November, where he met
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as well as observations of the life-history and breeding habits of many tropical frogs and fishes.
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themselves, which it has been my privilege to name and describe." He lists new species as follows:
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680:(Sitwell and Silva) had been ambushed at a conference and murdered, together with their policemen.
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818:, reaching it on 10 October. There he found a fisherman who had been fishing for Mr Loate of the
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in the wild. It took Budgett four African expeditions but in the end he succeeded in doing so.
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on 22 September. There he paid off the safari and spent a few days fishing unsuccessfully for
532:, lungfish and related groups could help to establish which group of fishes had given rise to
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and he had collected information about the fauna of the Gambia. Three papers were published.
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to house those extinct and extant animals that possessed lungs and fleshy pectoral fins with
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1239:(1946).The early evolution of fishes. Quarterly Review of Biology 21: 33–69, cited by Hall.
1216:(1907a).The development of Polypterus senegalus Cuv. Pages 195–290 in Kerr, J.G., ed. 1907b
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failed in his main goal he had gained experience; he had established the breeding period of
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During his subsequent study at Cambridge, Kerr had developed an ambition to make a study of
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However, the long periods spent in swampy, mosquito-ridden places left him debilitated with
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In 1890 Kerr had travelled on an expedition mounted by the Argentinean Navy to survey the
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He continued to work on his fishes in Cambridge but yearned to make another attempt on
1184:
Bartsch P., Gemballa S., Piotrowski T. (1997) The embryonic and larval development of
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as a crossopterygian and it is so described in many a classroom, although students of
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On his return to Cambridge, Budgett had to catch up on his reading for Part II of the
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Budgett, J H. (1907). Note on habits of Polypterus. Pages 291–292 in Kerr J G, ed.
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Budgett needed to find a position and applied for the post of Secretary to the
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Hall, B. K. (2001) John Samuel Budgett (1872–1904): In Pursuit of Polypterus,
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embryos. Harrington and Reid Hunt, a tutor in physiology at Columbia, reached
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Papers by Patterson (1982) and Noack (and fellow-authors, 1996) cited by Hall
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could help to answer questions about evolution. Perhaps the embryology of
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374:. He collected many samples and made drawings and notes. He discovered a
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Budgett and J. Graham Kerr. Cambridge (UK): Cambridge University Press.
822:. He engaged him and travelled back to Fashoda, arriving on 24 October.
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and his Cambridge students had shown in the 1870s and early 1880s that
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In March Budgett was elected to the Balfour Studentship . Named after
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on 27 July. There he did some fishing, catching many species but few
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On 29 August he gave up and proceeded further north by boat, passing
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1371:. Confirmation of William’s ownership can be found on their website
1141:. Proceedings of the Cambridge Philosophical Society 10s: 236–240.
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north to the Bolivian border. He had conducted original research on
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508:. He only had access to preserved specimens so his suggestion that
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1176:. Proceedings of the Zoological Society of London (1903): 25–26.
854:. Shipley was impressed by his friend’s knowledge and experience.
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analysis using both morphological and molecular data affirm that
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despite the health and safety risks. Dr Ansorge had brought back
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1166:. Transactions of the Zoological Society of London 16: 315–338.
1159:. Transactions of the Zoological Society of London 16: 115–134.
1148:. Transactions of the Zoological Society of London 15: 323–338.
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to the American Morphological Society. Harrington thought that
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All these papers were republished in Kerr, J.G., ed. (1907b).
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as an undergraduate and found that it provided a link between
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have realised the falsity of this position for many years...
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William was a keen microscopist and served on the council of
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was only a theory. Work in the field was needed to test it.
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1245:(1907). Biographical sketch. Pages 1–55 in Kerr J. G, ed.
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Budgett J. S. (1901a) On some points in the anatomy of
899:, the most westerly of the many streams into which the
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in June. A second object was added: to capture a live
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fry although he did catch fry of many other species.
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Budgett J. S. (1902) On the structure of the larval
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on 7 July. On 14 July they reached the mouth of the
1367:Post Code: BS9 1JP. This is now (2012) the home of
701:but failed to obtain the fertilised ova he wanted.
477:Periods. At that time little was known about them.
360:and this was the prime goal of the new expedition.
286:Budgett was educated first at a kindergarten, then
210:Budgett’s work remained the basis of understanding
1058:British Association for the Advancement of Science
492:. He had established a separate group (the tribe
1450:
834:Search for a Permanent Post and Final Expedition
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363:Budgett’s principal study on the expedition was
247:John Samuel Budgett was born at Redlands House,
1172:Budgett J. S. (1903b) Note on the spiracles of
1072:from the ancestral line leading to amphibians.
591:he observed and collected other fish. He found
574:
152:(16 June 1872 – 19 January 1904) was a British
1210:, Vol. 51, No. 5 (May 2001), pp. 399–407
1056:In a report to the 1907 annual meeting of the
16:British zoologist and embryologist (1872–1904)
755:. There it took about nine days to arrange a
563:At the time nothing was known about juvenile
408:. A third species has since been discovered.
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160:. He spent most of his short career on the
1115:
814:arriving on 5 October. Then he went on to
217:In an earlier expedition with Kerr to the
31:
1402:
1400:
1078:has no affinity with crossopterygians...
626:Anglo-Egyptian campaign against the Mahdi
1442:Bartsch and others (1997), cited by Hall
207:to interpret them and write the report.
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177:. Forty years after the publication of
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1133:Budgett J. S. (1900) Observations on
400:and subsequently published a paper.
1469:Alumni of Trinity College, Cambridge
1155:, and a description of the larva of
1104:is a living stem actinoptererygian.
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1385:
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624:on 26 May 1898 in the midst of the
449:, in Africa which were then called
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907:where he caught stern-wheeler for
744:but in the end this was given up.
683:Budgett traced the development of
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214:development for almost 100 years.
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404:was later given the common name
1484:19th-century British zoologists
1031:and the demonstration that the
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1064:amassed all the evidence that
876:. He wrote to him asking if a
779:). However he failed to catch
729:and the neighbouring streams.
567:. This was partly because its
556:, on the one hand, and to the
1:
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982:stages of the African fishes
713:In 1902 Budgett met the Rev.
671:Second Expedition: The Gambia
647:, about 650 km south of
579:Budgett thought first of the
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422:
242:
1125:Phyllomedusa hypochondrialis
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903:flows. He started first for
891:on 27 June 1903 and reached
810:. Then he took a steamer to
616:had already set out to find
575:First Expedition: The Gambia
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201:Zoological Society of London
7:
463:is now more usually called
292:University College, Bristol
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304:Trinity College, Cambridge
82:Trinity College, Cambridge
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840:London Zoological Society
751:with the Commissioner of
717:who was then resident in
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1382:, accessed 16 April 2012
1369:Trinity College, Bristol
1075:Romer (1946) asserted, "
734:Francis Maitland Balfour
1474:Scientists from Bristol
1358:Shipley in Kerr (1907b)
1116:Publications by Budgett
915:where he was told that
880:ship could take him to
583:but a friend suggested
429:Natural Sciences Tripos
1378:2 October 2013 at the
1062:Edwin Stephen Goodrich
484:had created an order:
397:Lepidobatrachus laevis
268:William Kitchen Parker
233:. The genus was named
230:Lepidobatrachus laevis
130:William Kitchen Parker
1489:British embryologists
1153:Protopterus annectens
747:Budgett travelled to
391:Lepidobatrachus asper
264:H.H. & S. Budgett
224:Lepidobatrachus asper
1186:Polypterus senegalus
1013:Synodontis ocellifer
998:Marcusenius budgetti
761:Polypterus senegalus
692:Polypterus senegalus
686:Gymnarchus niloticus
339:and the rest of the
120:Cambridge University
1479:Deaths from malaria
1424:Romer A. S. (1946).
1157:Polypterus lapradei
846:, Leipzig, Berlin,
753:British East Africa
698:Polypterus lapradei
614:Columbia University
496:) within the order
482:Thomas Henry Huxley
150:John Samuel Budgett
37:John Samuel Budgett
25:John Samuel Budgett
1180:Other publications
318:of South America.
1003:Gnathonemus gilli
771:. He caught many
608:Nathan Harrington
536:and hence to all
182:Origin of Species
147:
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126:Academic advisors
89:Scientific career
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1035:fin is really a
1008:Clarias budgetti
975:George Boulenger
964:blackwater fever
923:. He stopped at
911:. He stopped at
865:larvae from the
850:, Amsterdam and
378:new to science:
302:John went up to
205:John Graham Kerr
197:blackwater fever
138:John Graham Kerr
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1380:Wayback Machine
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707:
673:
661:Second Boer War
657:
655:Second Boer War
610:
577:
560:on the other".
522:Francis Balfour
443:
425:
402:Lepidobatrachus
381:Lepidobatrachus
323:Pilcomayo River
300:
288:Clifton College
245:
239:in his honour.
195:and he died of
111:
78:Alma mater
62:
61:19 January 1904
48:
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38:
26:
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12:
11:
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1396:
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1253:
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1247:
1243:Shipley, A. E.
1181:
1178:
1117:
1114:
1088:fish evolution
1053:
1049:Later Work on
1047:
1040:archipterygium
979:British Museum
971:
968:
943:
940:
897:Forcados River
874:Arthur Balfour
835:
832:
828:Lord Kitchener
820:British Museum
706:
703:
672:
669:
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609:
606:
576:
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518:Crossopterygii
498:Crossopterygii
486:Crossopterygii
442:
437:
424:
421:
416:Arthur Shipley
406:Budgett's Frog
311:at Cambridge.
299:
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260:Bristol Museum
244:
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236:Budgett's Frog
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134:Arthur Shipley
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727:Albert Nyanza
724:
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678:Commissioners
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490:lepidotrichia
487:
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466:Erpetoichthys
462:
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60:
56:
45:
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34:
29:
22:
19:
1438:
1429:
1420:
1411:
1394:Kerr (1907b)
1363:
1241:
1235:
1231:Patterson C.
1229:
1225:
1222:
1218:
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1205:
1199:
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1106:
1101:
1098:Phylogenetic
1096:
1091:
1083:
1079:
1076:
1074:
1069:
1065:
1055:
1050:
1044:
1028:
1025:urinogenital
1021:
1017:
1016:
1012:
1011:
1007:
1006:
1002:
1001:
997:
996:
993:
987:
983:
973:
959:
945:
936:P. senegalus
935:
933:
928:
916:
893:Sierra Leone
886:
882:Sierra Leone
871:
862:
858:
856:
837:
823:
807:
785:
780:
776:
772:
760:
746:
731:
722:
712:
708:
696:
690:
684:
682:
674:
658:
645:Atbara River
642:
637:
633:
629:
617:
611:
601:
592:
588:
578:
564:
562:
542:
529:
517:
516:belonged to
514:Calamichthys
513:
509:
506:Calamichthys
505:
501:
497:
494:Polypterinus
493:
479:
464:
461:Calamichthys
460:
457:Calamichthys
456:
450:
444:
439:
433:
426:
414:
410:
401:
395:
389:
379:
362:
357:
355:
344:
330:
327:Paraná River
320:
313:
301:
285:
276:
272:
257:
253:Stoke Bishop
246:
234:
228:
222:
216:
211:
209:
190:
185:
164:
158:embryologist
149:
148:
132:
116:Institutions
88:
63:(1904-01-19)
53:16 June 1872
18:
1464:1904 deaths
1459:1872 births
1282:Hall (2001)
1237:Romer, A S.
1139:Protopterus
988:Protopterus
767:, near the
715:John Roscoe
595:containing
372:Batrachians
358:Lepidosiren
346:Lepidosiren
337:cephalopods
280:Bristol Zoo
255:, Bristol.
70:Nationality
1453:Categories
1249:References
1214:Kerr, J.G.
1208:BioScience
1174:Polypterus
1164:Polypterus
1146:Polypterus
1135:Polypterus
1110:Polypterus
1102:Polypterus
1092:Polypterus
1084:Polypterus
1080:Polypterus
1070:Polypterus
1066:Polypterus
1051:Polypterus
1029:Polypterus
1027:system of
984:Polypterus
960:Polypterus
952:London Zoo
929:Polypterus
917:Polypterus
878:Royal Navy
863:Polypterus
859:Polypterus
824:Polypterus
808:Polypterus
781:Polypterus
773:Polypterus
723:Polypterus
638:Polypterus
634:Polypterus
630:Polypterus
618:Polypterus
602:Polypterus
593:Polypterus
589:Polypterus
585:the Gambia
565:Polypterus
546:Teleostome
544:primitive
534:amphibians
530:Polypterus
526:embryology
510:Polypterus
502:Polypterus
471:Palaeozoic
452:Polypterus
440:Polypterus
423:Graduation
368:Amphibians
316:Gran Chaco
243:Early life
219:Gran Chaco
212:Polypterus
186:Polypterus
166:Polypterus
107:Embryology
49:1872-06-16
1037:uniserial
942:Last Days
889:Liverpool
844:Frankfurt
804:Gondokoro
538:tetrapods
384:with two
325:from the
309:Pitt Club
298:Cambridge
175:amphibian
154:zoologist
1376:Archived
1195:Dean, B.
948:formalin
887:He left
816:Khartoum
725:ova was
649:Khartoum
554:Dipnoans
480:In 1861
475:Mesozoic
341:molluscs
332:Nautilus
179:Darwin’s
977:of the
956:Clifton
852:Antwerp
848:Hanover
812:Fashoda
788:Wadelai
749:Entebbe
738:Mombasa
695:and 36
665:malaria
569:habitat
558:Ganoids
548:to the
386:species
365:Anurous
249:Bristol
193:malaria
102:Zoology
73:British
970:Legacy
925:Muriji
921:Dakmon
909:Lokoja
905:Burutu
792:Nimule
757:Safari
719:Uganda
550:sharks
447:genera
351:Amazon
171:bichir
95:Fields
994:FISH
901:Niger
867:Niger
765:Fajao
742:okapi
622:Cairo
376:genus
162:genus
1373:here
1137:and
986:and
913:Assé
800:Legu
796:Kiri
581:Nile
552:and
512:and
504:and
500:for
473:and
455:and
394:and
227:and
156:and
58:Died
43:Born
777:sud
597:ova
370:or
1455::
1399:^
1387:^
1287:^
1257:^
1060:,
798:,
794:,
790:,
540:.
459:.
388:,
169:(
51:)
47:(
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