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Paddlefish (family Polyodontidae) are a family of
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
belonging to order
Acipenseriformes Acipenseriformes is an order (biology), order of basal (phylogenetics), basal Actinopterygii, ray-finned fishes that includes living and fossil sturgeons and paddlefishes (Acipenseroidei), as well as the extinct family (biology), families C ...
, and one of two living groups of the order alongside
sturgeon Sturgeon (from Old English ultimately from Proto-Indo-European language, Proto-Indo-European *''str̥(Hx)yón''-) is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the ...
s (Acipenseridae). They are distinguished from other fish by their elongated
rostra The Rostra () was a large platform built in the city of Rome that stood during the republican and imperial periods. Speakers would stand on the rostra and face the north side of the Comitium towards the senate house and deliver orations to t ...
, which are thought to enhance
electroreception Electroreception and electrogenesis are the closely related biological abilities to perceive electrical stimuli and to generate electric fields. Both are used to locate prey; stronger electric discharges are used in a few groups of fishes, such ...
to detect prey. Paddlefish have been referred to as " primitive fish" because the Acipenseriformes are among the earliest diverging lineages of ray-finned fish, having diverged from all other living groups over 300 million years ago. Both living and fossil paddlefish are found almost exclusively in North America and China. Eight species are known: Six of those species are extinct, and known only from fossils (five from North America, one from China),
one of the
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
species, the
American paddlefish The American paddlefish (''Polyodon spathula''), also known as a Mississippi paddlefish, spoon-billed cat, or spoonbill, is a species of ray-finned fish. It is the last extant taxon, living species of paddlefish (Polyodontidae). This family is ...
(''Polyodon spathula''), is native to the
Mississippi River The Mississippi River is the main stem, primary river of the largest drainage basin in the United States. It is the second-longest river in the United States, behind only the Missouri River, Missouri. From its traditional source of Lake Ita ...
basin in the U.S. The other is the Chinese paddlefish (''Psephurus gladius''), which was declared extinct in 2022 following a 2019 recommendation; the species has not been sighted in the Yangtze River Basin in China since 2003. Chinese paddlefish are also commonly referred to as "Chinese swordfish", or "elephant fish". The earliest known paddlefish is '' Protopsephurus'', from the
early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
(
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
) of China, dating to around 120 million years ago. Paddlefish populations have declined dramatically throughout their historic range as a result of
overfishing Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing Fish stocks, fish stock), resu ...
, pollution, and the encroachment of human development, including the construction of dams that have blocked their seasonal upward migration to ancestral spawning grounds. Other detrimental effects include alterations of rivers which have changed natural flows resulting in the loss of spawning habitat and nursery areas.


Morphology

Paddlefish as a group are one of the few organisms that retain a
notochord The notochord is an elastic, rod-like structure found in chordates. In vertebrates the notochord is an embryonic structure that disintegrates, as the vertebrae develop, to become the nucleus pulposus in the intervertebral discs of the verteb ...
past the embryonic stage. Paddlefish have very few
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
s and their bodies mostly consist of cartilage with the notochord functioning as a soft spine. During the initial stages of development from embryo to fry, paddlefish have no rostrum (snout). It begins to form shortly after hatching. The rostrum of the Chinese paddlefish was narrow and sword-like whereas the rostrum of the American paddlefish is broad and paddle-like. Some common morphological characteristics of paddlefish include a spindle-shaped, smooth-skinned scaleless body, heterocercal tail, and small poorly developed eyes. Unlike the filter-feeding American paddlefish, Chinese paddlefish were
piscivores A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
, and highly predatory. Their jaws were more forward pointing which suggested they foraged primarily on small fishes in the water column, and occasionally on shrimp,
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
fishes, and crabs. The jaws of the American paddlefish are distinctly adapted for filter feeding only. They are ram suspension filter feeders with a diet that consists primarily of
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
, and occasionally small insects, insect larvae, and small fish. The largest Chinese paddlefish on record measured in length, and was estimated to weigh a few thousand pounds. They commonly reached and . Although the American paddlefish is one of the largest freshwater fishes in North America, their recorded lengths and weights fell short in comparison to the larger Chinese paddlefish. American paddlefish commonly reach or more in length and can weigh more than . The largest American paddlefish on record was caught in 1916 in Okoboji Lake, Iowa. The fish was taken with a spear, and measured long and in the girth. A report published by J.R. Harlan and E.B. Speaker (1969) in ''Iowa Fish and Fishing'' states that the fish weighed over . The world record paddlefish caught on rod and reel weighed and was long. The fish was caught by Clinton Boldridge in a 5 acre pond in Atchison County, Kansas on 5 May 2004. However, the record would be broken an additional two times in 2020: On 28 June 2020, an Oklahoma man caught a 146 pound paddlefish in Keystone Lake, west of
Tulsa Tulsa ( ) is the second-most-populous city in the state of Oklahoma, after Oklahoma City, and the 48th-most-populous city in the United States. The population was 413,066 as of the 2020 census. It is the principal municipality of the Tul ...
. Later on 23 July 2020, the record was broken again when another Oklahoma man caught a 151 pound, nearly 6 foot long paddlefish in the same lake. Scientists once believed paddlefish used their rostrums to excavate bottom substrate, but have since determined with the aid of
electron microscopy An electron microscope is a microscope that uses a beam of electrons as a source of illumination. It uses electron optics that are analogous to the glass lenses of an optical light microscope to control the electron beam, for instance focusing i ...
that paddlefish rostrums are covered in electroreceptors called ampullae. These ampullae are densely packed within star-shaped bone projections that branch out from the rostrum. The electroreceptors can detect weak electrical fields which not only signal the presence of prey items in the water column, such as
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
which is the primary diet of the American paddlefish, but they can also detect the individual feeding and swimming movements of zooplankton's
appendages An appendage (or outgrowth) is an external body part or natural prolongation that protrudes from an organism's body such as an arm or a leg. Protrusions from single-celled bacteria and archaea are known as cell-surface appendages or surface app ...
. Paddlefish have poorly developed eyes, and rely on their electroreceptors for foraging. However, the rostrum is not the paddlefish's sole means of food detection. Some reports incorrectly suggest that a damaged rostrum would render paddlefish less capable of foraging efficiently to maintain good health. Laboratory experiments, and field research indicate otherwise. In addition to electroreceptors on the rostrum, paddlefish also have sensory pores covering nearly half of the skin surface extending from the rostrum to the top of the head down to the tips of the operculum (gill flaps). Paddlefish with damaged or abbreviated rostrums are still able to forage adequately.


Habitat and historic range

Over the past half century, paddlefish populations have been on the decline. Attributable causes are overfishing, pollution, and the encroachment of human development, including the construction of dams which block their seasonal upward migration to ancestral spawning grounds. Other detrimental effects include alterations of rivers which have changed the natural flow, and resulted in the loss of spawning habitat and nursery areas. American paddlefish have been extirpated from much of their Northern peripheral range, including the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes spanning the Canada–United States border. The five lakes are Lake Superior, Superior, Lake Michigan, Michigan, Lake Huron, H ...
and Canada, New York,
Maryland Maryland ( ) is a U.S. state, state in the Mid-Atlantic (United States), Mid-Atlantic region of the United States. It borders the states of Virginia to its south, West Virginia to its west, Pennsylvania to its north, and Delaware to its east ...
and
Pennsylvania Pennsylvania, officially the Commonwealth of Pennsylvania, is a U.S. state, state spanning the Mid-Atlantic (United States), Mid-Atlantic, Northeastern United States, Northeastern, Appalachian, and Great Lakes region, Great Lakes regions o ...
. There is growing concern about their populations in other states. The Chinese paddlefish was considered anadromous with upstream migration, however little is known about their migration habits and population structure. They were endemic to the Yangtze River Basin in China where they lived primarily in the broad surfaced main stem rivers and shoal zones along the
East China Sea The East China Sea is a marginal sea of the Western Pacific Ocean, located directly offshore from East China. China names the body of water along its eastern coast as "East Sea" (, ) due to direction, the name of "East China Sea" is otherwise ...
. Research suggests they preferred to navigate the middle and lower layers of the water column, and occasionally swam into large lakes. There have been no sightings of Chinese paddlefish since 2003, and were declared extinct in 2019. Past attempts of
artificial propagation Plant propagation is the process by which new plants grow from various sources, including seeds, cuttings, and other plant parts. Plant propagation can refer to both man-made and natural processes. Propagation typically occurs as a step in the o ...
for restoration purposes failed because of difficulties encountered in keeping captive fish alive. American paddlefish are native to the Mississippi River basin from New York to Montana and south to the Gulf of Mexico. They have been found in several Gulf Slope drainages in medium to large rivers with long, deep sluggish pools, as well as in backwater lakes and bayous. In Texas, paddlefish occurred historically in the Angelina River, Big Cypress Bayou, Neches River, tributaries of the Red River, Sabine River, San Jacinto River, Sulphur River, and Trinity River. Their historical range also included occurrences in Canada in
Lake Huron Lake Huron ( ) is one of the five Great Lakes of North America. It is shared on the north and east by the Canadian province of Ontario and on the south and west by the U.S. state of Michigan. The name of the lake is derived from early French ex ...
and Lake Helen, and in 26~27 other states in the United States. The Ontario Ministry of Natural Resources listed the paddlefish as extirpated from
Ontario, Canada Ontario is the southernmost Provinces and territories of Canada, province of Canada. Located in Central Canada, Ontario is the Population of Canada by province and territory, country's most populous province. As of the 2021 Canadian census, it ...
under their Endangered Species Act. The
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological ...
lists the Canadian populations of paddlefish as extirpated, noting there have been no Canadian records since the early 1900s and distribution in Canada was highly peripheral. As a species, the American paddlefish is classified as vulnerable (VU) on the IUCN Red List, and its international trade has been restricted since June 1992 under Appendix II of the Convention of International Trade in Endangered Species of Wild Flora and Fauna (
CITES CITES (shorter acronym for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of inte ...
).


Life cycle

Paddlefish are long-lived, and sexually late maturing. Females do not begin spawning until they are six to twelve years old, some even as late as sixteen to eighteen years old. Males begin spawning around age four to seven, some as late as nine or ten years of age. Paddlefish spawn in late spring if the proper combination of events occur; these include water flow, temperature, photoperiod, and availability of gravel substrates suitable for spawning. If all the conditions are not met, paddlefish do not spawn. Research suggests females do not spawn every year, rather they spawn every second or third year while males spawn more frequently, typically every year or every other year. Paddlefish migrate upstream to spawn, and prefer silt-free gravel bars that would otherwise be exposed to air, or covered by very shallow water were it not for the rises in the river from snow melt and annual spring rains that cause flooding. They are broadcast spawners, also referred to as mass spawners or synchronous spawners. Fertilization occurs externally:
Gravid In biology and medicine, gravidity and parity are the number of times a female has been pregnant (gravidity) and carried the pregnancies to a viable gestational age (parity). These two terms are usually coupled, sometimes with additional terms, t ...
females release their eggs into the water over bare rocks or gravel at the same time males release their sperm. The eggs are adhesive and stick to the rocky substrate. The young are swept downstream after hatching and grow to adulthood in deep freshwater pools.


Propagation and culture

The advancements in biotechnology in paddlefish propagation and rearing of captive stock indicate significant improvements in reproduction success, adaptation and survival rates of paddlefish cultured for broodstock development and stock rehabilitation. Such improvements have led to successful practices in reservoir ranching and pond rearing, creating an increasing interest in the global market for paddlefish polyculture. In a cooperative scientific effort in the early 1970s between the U.S. Fish & Wildlife Service and its former
USSR The Union of Soviet Socialist Republics. (USSR), commonly known as the Soviet Union, was a List of former transcontinental countries#Since 1700, transcontinental country that spanned much of Eurasia from 1922 until Dissolution of the Soviet ...
counterpart, American paddlefish were imported into the former USSR for
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. Nelu ...
, beginning with five-thousand hatched larvae from
Missouri Missouri (''see #Etymology and pronunciation, pronunciation'') is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. Ranking List of U.S. states and territories by area, 21st in land area, it border ...
hatcheries in the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. They were introduced into several rivers in Europe and Asia, and provided the first brood stock that were successfully reproduced in 1984–1986 in Russia. Paddlefish are now being raised in
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
,
Austria Austria, formally the Republic of Austria, is a landlocked country in Central Europe, lying in the Eastern Alps. It is a federation of nine Federal states of Austria, states, of which the capital Vienna is the List of largest cities in Aust ...
, the
Czech Republic The Czech Republic, also known as Czechia, and historically known as Bohemia, is a landlocked country in Central Europe. The country is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the south ...
, and the
Plovdiv Plovdiv (, ) is the List of cities and towns in Bulgaria, second-largest city in Bulgaria, 144 km (93 miles) southeast of the capital Sofia. It had a population of 490,983 and 675,000 in the greater metropolitan area. Plovdiv is a cultural hub ...
and Vidin regions in
Bulgaria Bulgaria, officially the Republic of Bulgaria, is a country in Southeast Europe. It is situated on the eastern portion of the Balkans directly south of the Danube river and west of the Black Sea. Bulgaria is bordered by Greece and Turkey t ...
. Reproduction was successful in 1988 and 1989, and resulted in the exportation of juvenile paddlefish to
Romania Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
and
Hungary Hungary is a landlocked country in Central Europe. Spanning much of the Pannonian Basin, Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia and ...
. In May 2006, specimens of different sizes and weights were caught by professional fisherman near Prahovo in the
Serbia , image_flag = Flag of Serbia.svg , national_motto = , image_coat = Coat of arms of Serbia.svg , national_anthem = () , image_map = , map_caption = Location of Serbia (gree ...
n part of the
Danube River The Danube ( ; see also other names) is the second-longest river in Europe, after the Volga in Russia. It flows through Central and Southeastern Europe, from the Black Forest south into the Black Sea. A large and historically important riv ...
. In 1988, fertilized paddlefish eggs and larvae from Missouri hatcheries were first introduced into China. Since that time, China imports approximately 4.5 million fertilized eggs and larvae every year from hatcheries in Russia, and the United States. Some of the paddlefish are polycultured in carp ponds, and sold to restaurants while others are cultured for brood stock and caviar production. China has also exported paddlefish to Cuba, where they are farmed for caviar production.


Classification

There is one currently
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
genus in this family, one recently
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
and five extinct genera known exclusively from
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s. Classification following , with '' Parapsephurus'' and '' Pugiopsephurus'' added in : * genus '' Protopsephurus'' Lu, 1994 (Early Cretaceous, China) ** species ''Protopsephurus liui'' Lu, 1994 * genus '' Pugiopsephurus'' Hilton ''et al.'', 2023 (Late Cretaceous, North America) (''
Incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'') ** species ''Pugiopsephurus inundatus'' Hilton ''et al.'', 2023 * clade Polyodonti ** genus '' Paleopsephurus'' MacAlpin, 1947 (Late Cretaceous, North America) *** species ''Paleopsephurus wilsoni'' MacAlpin, 1947 ** genus '' Parapsephurus'' Hilton ''et al.'', 2023 (Late Cretaceous, North America) *** species ''Parapsephurus willybemisi'' Hilton ''et al.'', 2023 ** subfamily Polyodontinae *** genus '' Psephurus'' Günther, 1873 **** ''Psephurus gladius'' E. von Martens, 1862 Chinese paddlefish (extinct c. 2003) *** tribe Polyodontini **** genus '' Crossopholis''
Cope A cope ( ("rain coat") or ("cape")) is a liturgical long mantle or cloak, open at the front and fastened at the breast with a band or clasp. It may be of any liturgical colour. A cope may be worn by any rank of the Catholic or Anglican clerg ...
'','' 1883
(
Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
, North America) ***** species ''Crossopholis magnicaudatus'' Cope, 1883 **** genus '' Polyodon'' Lacépède, 1797 (
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
-Recent, North America) ***** '' Polyodon spathula'' Walbaum, 1792
American paddlefish The American paddlefish (''Polyodon spathula''), also known as a Mississippi paddlefish, spoon-billed cat, or spoonbill, is a species of ray-finned fish. It is the last extant taxon, living species of paddlefish (Polyodontidae). This family is ...
***** ' Grande & Bemis, 1991 Relationships of the genera are from .


Notes


References


External links


One hour PBS documentary


– containing many photographs of ''Psepherus''.
images and movies of the paddlefish ''(Polyodon spathula)'' ARKive

FishBase entry for Polyodontidae



Fisheries.org

Paddlefish Fisheries Management

Stochastic synchronization of electroreceptors in the paddlefish
{{Authority control Polyodontidae Acipenseriformes Freshwater fish of North America Taxa named by Charles Lucien Bonaparte Extant Barremian first appearances