John Samuel Budgett
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John Samuel Budgett (16 June 1872 – 19 January 1904) was a British
zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and d ...
and embryologist. He spent most of his short career on the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''
Polypterus ''Polypterus'' is a genus of freshwater fish in the bichir family (Polypteridae) of order Polypteriformes. The type species is the Nile bichir (''P. bichir''). Fish in this genus live in various areas in Africa. ''Polypterus'' is the only kno ...
'' (
bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pu ...
). 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 amphibian. Forty years after the publication of Darwin’s
Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
, 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 ''Polypterus'' in the wild. It took Budgett four African expeditions but in the end he succeeded in doing so. However, the long periods spent in swampy, mosquito-ridden places left him debilitated with
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. S ...
and he died of blackwater fever 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
Zoological Society of London The Zoological Society of London (ZSL) is a charity devoted to the worldwide conservation of animals and their habitats. It was founded in 1826. Since 1828, it has maintained the London Zoo, and since 1931 Whipsnade Park. History On 29 ...
. 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
John Graham Kerr Sir John Graham Kerr (18 September 1869 – 21 April 1957), known to his friends as Graham Kerr, was a British embryologist and Unionist Member of Parliament (MP). He is best known for his studies of the embryology of lungfishes. He was involv ...
to interpret them and write the report.Shipley in Kerr (1907b) Budgett’s work remained the basis of understanding ''Polypterus'' development for almost 100 years.Hall (2001) In an earlier expedition with Kerr to the
Gran Chaco The Gran Chaco or Dry Chaco is a sparsely populated, hot and semiarid lowland natural region of the Río de la Plata basin, divided among eastern Bolivia, western Paraguay, northern Argentina, and a portion of the Brazilian states of Mato ...
in central South America, Budgett did useful work on the amphibians of that area and discovered two species of a new genus '' Lepidobatrachus asper'' and '' Lepidobatrachus laevis''. The genus was named '' Budgett's Frog'' in his honour.


Early life

John Samuel Budgett was born at Redlands House,
Bristol Bristol () is a city, ceremonial county and unitary authority in England. Situated on the River Avon, it is bordered by the ceremonial counties of Gloucestershire to the north and Somerset to the south. Bristol is the most populous city in ...
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,
Stoke Bishop Stoke Bishop is a medium-sized outer city suburb in the north-west of Bristol, located in between Westbury-on-Trym, Sneyd Park, and Sea Mills. Although relatively low, Stoke Bishop's population has increased due to substantial infilling on the ...
, Bristol. William was a keen microscopist and served on the council of
Bristol Museum Bristol Museum & Art Gallery is a large museum and art gallery in Bristol, England. The museum is situated in Clifton, about from the city centre. As part of Bristol Culture it is run by the Bristol City Council with no entrance fee. It holds ...
. He was a keen amateur student of zoology and encouraged his children to study it. Professionally he was a partner in H.H. & S. Budgett, 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
William Kitchen Parker William Kitchen Parker FRS FRMS (23 June 1823 – 3 July 1890) was a British physician, zoologist and comparative anatomist. From a humble beginning he became Hunterian Professor of Anatomy and Physiology in the College of Surgeons of Engl ...
. 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. 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. 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
Bristol Zoo Bristol Zoo was a zoo in the city of Bristol in South West England. The zoo's stated mission was to "maintain and defend” biodiversity through breeding endangered species, conserving threatened species and habitats and promoting a wider unders ...
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. Budgett was educated first at a kindergarten, then
Clifton College ''The spirit nourishes within'' , established = 160 years ago , closed = , type = Public schoolIndependent boarding and day school , religion = Christian , president = , head_label = Head of College , hea ...
. 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
University College, Bristol University College, Bristol was an educational institution which existed from 1876 to 1909. It was the predecessor institution to the University of Bristol, which gained a royal charter in 1909. During its time the college mainly served the midd ...
. There he received help in the formal study of zoology but much of his practical skill was self-taught.


Cambridge

John went up to
Trinity College, Cambridge Trinity College is a constituent college of the University of Cambridge. Founded in 1546 by King Henry VIII, Trinity is one of the largest Cambridge colleges, with the largest financial endowment of any college at either Cambridge or Oxford. ...
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
Pitt Club The University Pitt Club, popularly referred to as the Pitt Club, the UPC, or merely as Club, is a private members' club of the University of Cambridge, with a previously male-only membership but now open to both men and women. History The ...
at Cambridge.Kerr (1907b) 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 The Gran Chaco or Dry Chaco is a sparsely populated, hot and semiarid lowland natural region of the Río de la Plata basin, divided among eastern Bolivia, western Paraguay, northern Argentina, and a portion of the Brazilian states of Mato ...
of South America. In 1890 Kerr had travelled on an expedition mounted by the Argentinean Navy to survey the
Pilcomayo River Pilcomayo (in Hispanicized spelling) (Quechua Pillkumayu or Pillku Mayu, ''pillku'' red, ''mayu'' river, "red river", Guarani Ysyry Araguay ) is a river in central South America. At long, it is the longest western tributary of the Paraguay River ...
from the Paraná River north to the Bolivian border. He had conducted original research on ''
Nautilus The nautilus (, ) is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in ...
'' as an undergraduate and found that it provided a link between cephalopods and the rest of the molluscs. 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 ''
Lepidosiren The South American lungfish (''Lepidosiren paradoxa'') is the single species of lungfish found in swamps and slow-moving waters of the Amazon, Paraguay, and lower Paraná River basins in South America. Notable as an obligate air-breather, it ...
'' lived in the
Amazon Amazon most often refers to: * Amazons, a tribe of female warriors in Greek mythology * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon River, in South America * Amazon (company), an American multinational technolog ...
so when Kerr learned in 1890 that a soldier had caught an eel-like fish he wondered whether it might be a lungfish. During his subsequent study at Cambridge, Kerr had developed an ambition to make a study of ''Lepidosiren'' and this was the prime goal of the new expedition. Budgett’s principal study on the expedition was Anurous Amphibians or
Batrachian A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
s. He collected many samples and made drawings and notes. He discovered a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
new to science: ''
Lepidobatrachus ''Lepidobatrachus'' is a genus of ceratophryidid frogs. They are commonly known as Paraguay horned frogs or Budgett's frogs (in honor of John Samuel Budgett, who described the genus), although the latter technically describes a specific specie ...
'' with two
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, '' Lepidobatrachus asper'' and '' Lepidobatrachus laevis'' and subsequently published a paper. Budgett (1899a) ''Lepidobatrachus'' was later given the common name Budgett's Frog. A third species has since been discovered. 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 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.
Arthur Shipley Sir Arthur Everett Shipley GBE FRS (10 March 1861 – 22 September 1927) was an English zoologist and Vice-Chancellor of the University of Cambridge. Biography Shipley was born in Walton-on-Thames, Surrey on 10 March 1861. He was brought up in ...
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 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 udgett'sskill in manipulation and his peculiar knowledge of the use of reagents.".


Graduation

On his return to Cambridge, Budgett had to catch up on his reading for Part II of the
Natural Sciences Tripos The Natural Sciences Tripos (NST) is the framework within which most of the science at the University of Cambridge is taught. The tripos includes a wide range of Natural Sciences from physics, astronomy, and geoscience, to chemistry and biology, ...
(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. 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 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.


Polypterus

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 genera, in Africa which were then called ''
Polypterus ''Polypterus'' is a genus of freshwater fish in the bichir family (Polypteridae) of order Polypteriformes. The type species is the Nile bichir (''P. bichir''). Fish in this genus live in various areas in Africa. ''Polypterus'' is the only kno ...
'' and ''Calamichthys''. ''Calamichthys'' is now more usually called '' Erpetoichthys''. These are the sole survivors of a vast group which flourished in the Palaeozoic and
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
Periods. At that time little was known about them. In 1861
Thomas Henry Huxley Thomas Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist specialising in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The stori ...
had created an order: Crossopterygii to house those extinct and extant animals that possessed lungs and fleshy pectoral fins with
lepidotrichia Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as see ...
. He had established a separate group (the tribe ''Polypterinus'') within the order ''Crossopterygii'' for ''Polypterus'' and ''Calamichthys''. He only had access to preserved specimens so his suggestion that ''Polypterus'' and ''Calamichthys'' belonged to ''Crossopterygii'' was only a theory. Work in the field was needed to test it. Francis Balfour and his Cambridge students had shown in the 1870s and early 1880s that
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos ...
could help to answer questions about evolution. Perhaps the embryology of ''Polypterus'', lungfish and related groups could help to establish which group of fishes had given rise to amphibians and hence to all
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s. 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
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachi ...
s and Dipnoans, on the one hand, and to the
Ganoid A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as w ...
s on the other". At the time nothing was known about juvenile ''Polypterus''. This was partly because its
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
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


First Expedition: The Gambia

Budgett thought first of the
Nile The Nile, , Bohairic , lg, Kiira , Nobiin: Áman Dawū is a major north-flowing river in northeastern Africa. It flows into the Mediterranean Sea. The Nile is the longest river in Africa and has historically been considered the longest ...
but a friend suggested
the Gambia The Gambia,, ff, Gammbi, ar, غامبيا officially the Republic of The Gambia, is a country in West Africa. It is the smallest country within mainland AfricaHoare, Ben. (2002) ''The Kingfisher A-Z Encyclopedia'', Kingfisher Publicatio ...
. 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 ''Polypterus'' he observed and collected other fish. He found ''Polypterus'' containing
ova , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...
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 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. Three papers were published.


Nathan Harrington

Quite independently, Nathan Harrington (1870–1899), an experienced field biologist and a doctoral student at
Columbia University Columbia University (also known as Columbia, and officially as Columbia University in the City of New York) is a private research university in New York City. Established in 1754 as King's College on the grounds of Trinity Church in Manhatt ...
had already set out to find ''Polypterus'' embryos. Harrington and Reid Hunt, a tutor in physiology at Columbia, reached
Cairo Cairo ( ; ar, القاهرة, al-Qāhirah, ) is the Capital city, capital of Egypt and its largest city, home to 10 million people. It is also part of the List of urban agglomerations in Africa, largest urban agglomeration in Africa, List of ...
on 26 May 1898 in the midst of the Anglo-Egyptian campaign against the Mahdi. 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 ''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. 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 The Atbarah River ( ar, نهر عطبرة; transliterated: Nahr 'Atbarah), also known as the Black Nile, is a river in northeast Africa. It rises in northwest Ethiopia, approximately 50 km north of Lake Tana and 30 km west of Gond ...
, about 650 km south of
Khartoum Khartoum or Khartum ( ; ar, الخرطوم, Al-Khurṭūm, din, Kaartuɔ̈m) is the capital of Sudan. With a population of 5,274,321, its metropolitan area is the largest in Sudan. It is located at the confluence of the White Nile, flowing n ...
. 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.


Second Boer War

When Budgett returned to England in the Autumn of 1899, he found the country in a turmoil on the eve of the
Second Boer War The Second Boer War ( af, Tweede Vryheidsoorlog, , 11 October 189931 May 1902), also known as the Boer War, the Anglo–Boer War, or the South African War, was a conflict fought between the British Empire and the two Boer Republics (the South ...
. 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
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. S ...
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.


Second Expedition: The Gambia

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 Commissioners (Sitwell and Silva) had been ambushed at a conference and murdered, together with their policemen. Budgett traced the development of ''
Gymnarchus niloticus ''Gymnarchus niloticus'' – commonly known as the ''aba'', ''aba aba'', frankfish, freshwater rat-tail, ''poisson-cheval'', or African knifefish – is an electric fish, and the only species in the genus ''Gymnarchus'' and the family Gymnarchid ...
'' and made observations on other fish, frogs, mosquitoes and other insects. He returned in August. He had caught 127 ''
Polypterus senegalus ''Polypterus senegalus'', the Senegal bichir, gray bichir or Cuvier's bichir, and sometimes called the "dinosaur eel" (a misnomer, as the creature is neither an eel nor a dinosaur), "dinosaur bichir", or "dragon fish" is in the pet trade due to ...
'' and 36 '' Polypterus lapradei'' but failed to obtain the fertilised ova he wanted.


Assistant Curator and Third Expedition

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. In 1902 Budgett met the Rev.
John Roscoe John Roscoe (1861–1932) was an Anglican missionary to East Africa. He conducted anthropological data collection of the Africans he encountered on mission. Roscoe was born in 1861, during the height of the Victorian era. Roscoe's career heavi ...
who was then resident in
Uganda }), is a landlocked country in East Africa. The country is bordered to the east by Kenya, to the north by South Sudan, to the west by the Democratic Republic of the Congo, to the south-west by Rwanda, and to the south by Tanzania. The sou ...
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 ''Polypterus'' ova was Albert Nyanza and the neighbouring streams. In March Budgett was elected to the Balfour Studentship . Named after
Francis Maitland Balfour Francis (Frank) Maitland Balfour, known as F. M. Balfour, (10 November 1851 – 19 July 1882) was a British biologist. He lost his life while attempting the ascent of Mont Blanc. He was regarded by his colleagues as one of the greatest biologist ...
, 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
Mombasa Mombasa ( ; ) is a coastal city in southeastern Kenya along the Indian Ocean. It was the first capital of the British East Africa, before Nairobi was elevated to capital city status. It now serves as the capital of Mombasa County. The town is ...
in June. A second object was added: to capture a live
Okapi The okapi (; ''Okapia johnstoni''), also known as the forest giraffe, Congolese giraffe, or zebra giraffe, is an artiodactyl mammal that is endemic to the northeast Democratic Republic of the Congo in central Africa. It is the only species ...
but in the end this was given up. Budgett travelled to
Entebbe Entebbe is a city in Central Uganda. Located on a Lake Victoria peninsula, approximately southwest of the Ugandan capital city, Kampala. Entebbe was once the seat of government for the Protectorate of Uganda prior to independence, in 1962. T ...
with the Commissioner of
British East Africa East Africa Protectorate (also known as British East Africa) was an area in the African Great Lakes occupying roughly the same terrain as present-day Kenya from the Indian Ocean inland to the border with Uganda in the west. Controlled by Bri ...
. There it took about nine days to arrange a Safari. 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 ''Polypterus senegalus''. 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 Fajao, near the
Murchison Falls Murchison Falls, also known as Kabalega Falls, is a waterfall at the apex of Lake Albert on the Victoria Nile in Uganda. At the top of Murchison Falls, the Nile forces its way through a gap in the rocks, only wide, and tumbles , before flowing ...
. He caught many ''Polypterus'' females (locally called "Intontos") laying or having laid eggs. He was certain that the fry must swarm in the floating vegetation (''sud''). However he failed to catch ''Polypterus'' fry although he did catch fry of many other species. On 29 August he gave up and proceeded further north by boat, passing Wadelai,
Nimule Nimule is a city in the southern part of South Sudan in Magwi County, Eastern Equatoria. It lies approximately , by road, southeast of Juba, the capital of South Sudan and largest city in the country. The town also lies approximately , by road, no ...
, Kiri, Legu and reaching
Gondokoro Gondokoro island is located in Central Equatoria. The island was a trading-station on the east bank of the White Nile in Southern Sudan, south of Khartoum. Its importance lay in the fact that it was within a few kilometres of the limit of naviga ...
on 22 September. There he paid off the safari and spent a few days fishing unsuccessfully for ''Polypterus''. Then he took a steamer to
Fashoda Kodok or Kothok ( ar, كودوك), formerly known as Fashoda, is a town in the north-eastern South Sudanese state of Upper Nile State. Kodok is the capital of Shilluk country, formally known as the Shilluk Kingdom. Shilluk had been an independ ...
arriving on 5 October. Then he went on to
Khartoum Khartoum or Khartum ( ; ar, الخرطوم, Al-Khurṭūm, din, Kaartuɔ̈m) is the capital of Sudan. With a population of 5,274,321, its metropolitan area is the largest in Sudan. It is located at the confluence of the White Nile, flowing n ...
, reaching it on 10 October. There he found a fisherman who had been fishing for Mr Loate of the
British Museum The British Museum is a public museum dedicated to human history, art and culture located in the Bloomsbury area of London. Its permanent collection of eight million works is among the largest and most comprehensive in existence. It docum ...
. He engaged him and travelled back to Fashoda, arriving on 24 October. ''Polypterus'' were caught but they had already shed their eggs. He returned to Khartoum on 10 November, where he met Lord Kitchener. He was home by the end of the month.


Search for a Permanent Post and Final Expedition

Budgett needed to find a position and applied for the post of Secretary to the London Zoological Society. 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,
Frankfurt Frankfurt, officially Frankfurt am Main (; Hessian: , " Frank ford on the Main"), is the most populous city in the German state of Hesse. Its 791,000 inhabitants as of 2022 make it the fifth-most populous city in Germany. Located on it ...
, Leipzig, Berlin,
Hanover Hanover (; german: Hannover ; nds, Hannober) is the capital and largest city of the German state of Lower Saxony. Its 535,932 (2021) inhabitants make it the 13th-largest city in Germany as well as the fourth-largest city in Northern Germany ...
, Amsterdam and Antwerp. Shipley was impressed by his friend’s knowledge and experience. He continued to work on his fishes in Cambridge but yearned to make another attempt on ''Polypterus'' despite the health and safety risks. Dr Ansorge had brought back ''Polypterus'' larvae from the
Niger ) , official_languages = , languages_type = National languagesArthur Balfour Arthur James Balfour, 1st Earl of Balfour, (, ; 25 July 184819 March 1930), also known as Lord Balfour, was a British Conservative statesman who served as Prime Minister of the United Kingdom from 1902 to 1905. As foreign secretary in the ...
. He wrote to him asking if a
Royal Navy The Royal Navy (RN) is the United Kingdom's naval warfare force. Although warships were used by English and Scottish kings from the early medieval period, the first major maritime engagements were fought in the Hundred Years' War against ...
ship could take him to
Sierra Leone Sierra Leone,)]. officially the Republic of Sierra Leone, is a country on the southwest coast of West Africa. It is bordered by Liberia to the southeast and Guinea surrounds the northern half of the nation. Covering a total area of , Sierr ...
. The answer was negative but Balfour very generously paid for Budgett’s passage out of his own pocket. He left
Liverpool Liverpool is a city and metropolitan borough in Merseyside, England. With a population of in 2019, it is the 10th largest English district by population and its metropolitan area is the fifth largest in the United Kingdom, with a populat ...
on 27 June 1903 and reached
Sierra Leone Sierra Leone,)]. officially the Republic of Sierra Leone, is a country on the southwest coast of West Africa. It is bordered by Liberia to the southeast and Guinea surrounds the northern half of the nation. Covering a total area of , Sierr ...
on 7 July. On 14 July they reached the mouth of the Forcados River, the most westerly of the many streams into which the
Niger ) , official_languages = , languages_type = National languagesBurutu where he caught stern-wheeler for Lokoja. He stopped at Assé where he was told that ''Polypterus'' 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 Dakmon. He stopped at Muriji on 27 July. There he did some fishing, catching many species but few ''Polypterus''. 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. It was very wet but he worked hard, fishing and also collecting and interviewing local fishermen. His guide knew about ''P. Senegalus''. 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.


Last Days

On his return to England, Budgett was suffering bouts of malaria and his hands had sores from constantly handling
formalin Formaldehyde ( , ) ( systematic name methanal) is a naturally occurring organic compound with the formula and structure . The pure compound is a pungent, colourless gas that polymerises spontaneously into paraformaldehyde (refer to section F ...
. However he returned to Cambridge and began to work out his material. The post of Resident Superintendent at London Zoo 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 Clifton, returning to Cambridge early in January. By 9 January he had finished his drawings of the external features of the developing ''Polypterus'' ova. That evening he had an attack of blackwater fever. 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.


Legacy

George Boulenger George Albert Boulenger (19 October 1858 – 23 November 1937) was a Belgian-British zoologist who described and gave scientific names to over 2,000 new animal species, chiefly fish, reptiles, and amphibians. Boulenger was also an active botani ...
of the
British Museum The British Museum is a public museum dedicated to human history, art and culture located in the Bloomsbury area of London. Its permanent collection of eight million works is among the largest and most comprehensive in existence. It docum ...
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 stages of the African fishes ''Polypterus'' and ''Protopterus'' 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.... ndvery important collections of the fishes themselves, which it has been my privilege to name and describe." He lists new species as follows: FISH ''Marcusenius budgetti'' ''Gnathonemus gilli'' ''Clarias budgetti'' ''Synodontis ocellifer'' ''Rana budgetti'' 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 urinogenital system of ''Polypterus'' and the demonstration that the
crossopterygian Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includin ...
fin is really a uniserial archipterygium as well as observations of the life-history and breeding habits of many tropical frogs and fishes. As a memorial, Budgett’s friend Graham Kerr edited a book.Kerr (1907b) This contains all his papers as well as a biographical sketch by A. E. Shipley and assessments of his work by several authors


Later Work on ''Polypterus''

In a report to the 1907 annual meeting of the British Association for the Advancement of Science,
Edwin Stephen Goodrich Edwin Stephen Goodrich FRS (Weston-super-Mare, 21 June 1868 – Oxford, 6 January 1946), was an English zoologist, specialising in comparative anatomy, embryology, palaeontology, and evolution. He held the Linacre Chair of Zoology in the Univer ...
amassed all the evidence that ''Polypterus'' is not a crossopterygian, placing it within the palaeoniscids, the most primitive of the ray-finned (actinopterygian) fishes. This removed ''Polypterus'' from the ancestral line leading to amphibians. Romer (1946) asserted, "'' olypterus' has no affinity with crossopterygians... ''Polypterus'' 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 ''Polypterus'' as a crossopterygian and it is so described in many a classroom, although students of fish evolution have realised the falsity of this position for many years... ''Polypterus''... is not a crossopterygian, but an actinopterygian and hence can tell us nothing about crossopterygian anatomy and embryology."
Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis using both morphological and molecular data affirm that ''Polypterus'' is a living stem actinoptererygian. A 1997 studyBartsch and others (1997), cited by Hall 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 ''Polypterus'' and of urodele amphibians and other 'primitive' amphibians. Other questions that Budgett was hoping to answer are still unanswered.


Publications by Budgett

All these papers were republished in Kerr, J.G., ed. (1907b). 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. Also a description of a new genus. Quarterly Journal of Microscopical Sciences 2 : 305–333. 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. Budgett J. S. (1900) Observations on ''Polypterus'' and ''Protopterus''. Proceedings of the Cambridge Philosophical Society 10s: 236–240. Budgett J. S. (1901a) On some points in the anatomy of ''Polypterus''. Transactions of the Zoological Society of London 15: 323–338. 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''. Transactions of the Zoological Society of London 16: 115–134. Budgett J. S. (1902) On the structure of the larval ''Polypterus''. Transactions of the Zoological Society of London 16: 315–338. Budgett J. S. (1903a) Account of journey to Uganda. Proceedings of the Zoological Society of London: 2–10. Budgett J. S. (1903b) Note on the spiracles of ''Polypterus''. Proceedings of the Zoological Society of London (1903): 25–26.


Other publications

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. Acta Zoologica 78: 309–328, cited by Hall. Budgett, J H. (1907). Note on habits of Polypterus. Pages 291–292 in Kerr J G, ed. Dean, B. 1895. Fishes, Living and Fossil. An Outline of their Forms and Probable Relationships p. 149. Delhi (India): Narendra Publishing House., cited by Hall Goodrich, E. S. (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 Hall, B. K. (2001) John Samuel Budgett (1872–1904): In Pursuit of Polypterus
BioScience
Vol. 51, No. 5 (May 2001), pp. 399–407 Kerr, J.G. (1907a).The development of Polypterus senegalus Cuv. Pages 195–290 in Kerr, J.G., ed. 1907b 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 Budgett and J. Graham Kerr. Cambridge (UK): Cambridge University Press. Kerr, J.G. (1950). A Naturalist in the Gran Chaco. Cambridge (UK): Cambridge University Press, cited by Hall. 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 Patterson C. (1982). Morphology and interrelationships of primitive actinopterygian fishes. American Zoologist 22: 241–260, cited by Hall. Romer, A S. (1946).The early evolution of fishes. Quarterly Review of Biology 21: 33–69, cited by Hall. Shipley, A. E. (1907). Biographical sketch. Pages 1–55 in Kerr J. G, ed.


References

{{DEFAULTSORT:Budgett, John Samuel 1872 births 1904 deaths Alumni of Trinity College, Cambridge Scientists from Bristol Deaths from malaria 19th-century British zoologists British embryologists