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''Polypterus'' is 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 ...
of
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does incl ...
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
in the bichir
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
(
Polypteridae 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 ...
) of
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Polypteriformes. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specim ...
is the Nile bichir (''P. bichir''). Fish in this genus live in various areas in
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
. ''Polypterus'' is the only known
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
to have
lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of ...
s, but no
trachea The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air- breathing animals with lungs. The trachea extends from t ...
. The
etymology Etymology () The New Oxford Dictionary of English (1998) – p. 633 "Etymology /ˌɛtɪˈmɒlədʒi/ the study of the class in words and the way their meanings have changed throughout time". is the study of the history of the form of words ...
of the genus name derives from a combination of the Greek
prefix A prefix is an affix which is placed before the stem of a word. Adding it to the beginning of one word changes it into another word. For example, when the prefix ''un-'' is added to the word ''happy'', it creates the word ''unhappy''. Particul ...
πολυ-, ''poly-'' (many) and the root word πτερον, ''pteron'' (wing or fin) – "many fins".


Recoil aspiration

In shallow water, ''Polypterus'' inhales primarily through its spiracle (blowhole). Exhalation is powered by muscles in the torso. During exhalation, the bony scales in the upper chest become indented. When the muscles are relaxed, the bony scales spring back into position, generating negative pressure within the torso, resulting in a rapid intake of air through the spiracle. The air is nearly sufficient to fill the lungs. This is followed by one cycle of buccal (mouth) pumping, which "tops off" the lungs, with the surplus air from the buccal pumping process discharged through the
pharynx The pharynx (plural: pharynges) is the part of the throat behind the mouth and nasal cavity, and above the oesophagus and trachea (the tubes going down to the stomach and the lungs). It is found in vertebrates and invertebrates, though its st ...
. According to one hypothesis,
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
tetrapods may have inhaled in this way.


Discovery

''Polypterus'' was discovered, described, and named in 1802 by
Étienne Geoffroy Saint-Hilaire Étienne Geoffroy Saint-Hilaire (15 April 177219 June 1844) was a French naturalist who established the principle of "unity of composition". He was a colleague of Jean-Baptiste Lamarck and expanded and defended Lamarck's evolutionary theories ...
. It is 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 ...
of 10 green to yellow-brown species. Naturalists were unsure whether to regard it as a fish or an amphibian. If it were a fish, what type was it: bony, cartilaginous, or lungfish?Hall (2001) Some regarded ''Polypterus'' as a living fossil, part of the missing link between fishes and amphibians, helping to show how fish fins had evolved to become paired limbs. In 1861,
Thomas 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 ...
created the order
Crossopterygii 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 includi ...
to house animals, fossil and living, that possessed lungs and fleshy pectoral fins with lobes. He placed ''Polypterus'' and '' Calamoichthys'' within this order, allocating them to a new tribe, Polypterini, which he created especially for them. The weight of Huxley's authority allowed this allocation to last in textbooks and lectures long after it had been disproved. In the 1870s and 1880s, Francis Balfour and his students had shown 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 embr ...
could help to answer questions about the evolution of species. No one had studied the embryology of ''Polypterus''. Someone who could do this might prove the “missing link” theory and be greatly honoured, but it could be a dangerous quest. The only breeding ''Polypterus'' specimens were in swampy parts of African rivers. Africa was a turbulent place and swamps were rich breeding grounds for mosquitoes carrying
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. ...
. Two men, Nathan Harrington and
John Samuel Budgett John Samuel Budgett (16 June 1872 – 19 January 1904) was a British zoologist and embryologist. He spent most of his short career on the genus ''Polypterus'' (bichir). This is found in the lakes, river margins, swamps, and floodplains of tropi ...
, attempted to answer this question by making repeated expeditions to Africa. Harrington failed on his first attempt in 1898 and died early on his second in 1899 before he could reach his destination. Budgett failed in 1898/9, 1900, and 1902. He finally succeeded in 1903, but died of
blackwater fever Blackwater fever is a complication of malaria infection in which red blood cells burst in the bloodstream (hemolysis), releasing hemoglobin directly into the blood vessels and into the urine, frequently leading to kidney failure. The disease w ...
shortly after his return to England. He left excellent samples and drawings, but his only writing was a diary. Consequently, his results on ''Polypterus'' were written up and published by his friend
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 ...
. Drawing on this work, in 1907, E. S. Goodrich reported to the British Association the then current state of evidence 'against' ''Polypterus'' being a crossopterygian, placing it within the palaeoniscids, the most primitive actinopterygians. Much later, in 1946, Romer, confirmed this view, but he also wrote, "The weight of Huxley's 861opinion 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 realized 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." Hall (2001), relying on Patterson (1982) and Noack ''et al.'' (1996), writes, "Phylogenetic analyses using both morphological and molecular data affirm ''Polypterus'' as a living stem actinopterygian." Research is ongoing. Most of the conclusions drawn by Kerr from Budgett's specimens have been confirmed, but many questions remain. ''Polypterus'' has rarely been bred in captivity. The first success was that of ''Polypterus senegalus'' by Arnoult in 1964, a species spawned repeatedly since (see Hartl, 1981; Bartsch and Gemballa, 1992; Bartsch et al., 1997 and Schugardt, 1997). Shortly after Arnoult's success, a second species, ''Polypterus ornatipinnis'', was spawned by Armbrust for the first time (1966 and 1973) and bred subsequently by Azuma in 1986; Wolf, 1992; Bartsch and Britz, 1996. The third species successfully spawned in captivity was ''Polypterus endlicheri'' by Azuma in 1995. Zoo Basel have been successful in breeding ''Polypterus'' in captivity. In December 2005, several eggs were laid, and at the beginning of 2006, six young hatched. Within two months, they reached 10 cm (about 4 in). In 2014 researchers at McGill University (published in the journal Nature) turned to ''Polypterus'' to help show what might have happened when fish first attempted to walk out of the water. The team of researchers raised juvenile ''Polypterus'' on land for nearly a year, with the aim of revealing how these 'terrestrialized' fish looked and moved differently.


Species

* '' Polypterus ansorgii''
Boulenger Boulenger is a surname. Notable people with the surname include: * Benjamin Boulenger (born 1990), French footballer * Edward George Boulenger (1888–1946), British zoologist, director of aquarium at London Zoo * George Albert Boulenger (1858– ...
, 1910
(Guinean bichir) * '' Polypterus bichir'' Lacépède, 1803 (Nile bichir) ** ''P. b. lapradei''
Steindachner Franz Steindachner (11 November 1834 in Vienna – 10 December 1919 in Vienna) was an Austrian zoologist, ichthyologist, and herpetologist. He published over 200 papers on fishes and over 50 papers on reptiles and amphibians. Steindachner describ ...
, 1869
(junior synonym) * '' Polypterus congicus''
Boulenger Boulenger is a surname. Notable people with the surname include: * Benjamin Boulenger (born 1990), French footballer * Edward George Boulenger (1888–1946), British zoologist, director of aquarium at London Zoo * George Albert Boulenger (1858– ...
, 1898
(Congo bichir) * '' Polypterus delhezi''
Boulenger Boulenger is a surname. Notable people with the surname include: * Benjamin Boulenger (born 1990), French footballer * Edward George Boulenger (1888–1946), British zoologist, director of aquarium at London Zoo * George Albert Boulenger (1858– ...
, 1899
(Barred bichir) * '' Polypterus endlicherii'' Heckel, 1847 * '' Polypterus mokelembembe'' Schliewen &
Schäfer Schaefer is an alternative spelling and cognate for the German word ''schäfer'', meaning 'shepherd', which itself descends from the Old High German '' scāphare''. Variants "Shaefer", "Schäfer" (a standardized spelling in many German-speakin ...
, 2006
(Mokèlé-mbèmbé bichir) * '' Polypterus ornatipinnis''
Boulenger Boulenger is a surname. Notable people with the surname include: * Benjamin Boulenger (born 1990), French footballer * Edward George Boulenger (1888–1946), British zoologist, director of aquarium at London Zoo * George Albert Boulenger (1858– ...
, 1902
(Ornate bichir) * '' Polypterus palmas'' Ayres, 1850 *'' Polypterus polli'' J. P. Gosse, 1988 (Poll's bichir) * '' Polypterus retropinnis'' Vaillant, 1899 (West African bichir) * '' Polypterus senegalus''
Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in nat ...
, 1829
(Gray bichir) * '' Polypterus teugelsi'' Britz, 2004 (Cross River bichir) * '' Polypterus weeksii''
Boulenger Boulenger is a surname. Notable people with the surname include: * Benjamin Boulenger (born 1990), French footballer * Edward George Boulenger (1888–1946), British zoologist, director of aquarium at London Zoo * George Albert Boulenger (1858– ...
, 1898
(Mottled bichir)


References


Sources

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. Quoted 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. (1907), ''The development of Polypterus senegalus Cuv.'', Pages 195-290 in Kerr, J.G., (ed. 1907), ''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. 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 {{Taxonbar, from=Q1303295 Polypteridae Taxa named by Bernard Germain de Lacépède Freshwater fish genera