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Paddlefish (family Polyodontidae) are a family of
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
belonging to order
Acipenseriformes Acipenseriformes is an order of basal ray-finned fishes that includes living and fossil sturgeons and paddlefishes (Acipenseroidei), as well as the extinct families Chondrosteidae and Peipiaosteidae. They are the second earliest diver ...
, and one of two living groups of the order alongside sturgeons (Acipenseridae). They are distinguished from other fish by their titular elonglated rostrums, which are thought to enhance electroreception to detect prey. Paddlefish have been referred to as " primitive fish" because Acipenseriformes are amongst the earliest diverging lineages of ray-finned fish, having diverged from all other living groups over 300 million years ago. Paddlefish are almost exclusively North American and Chinese, both extant and in the fossil record. There are six known species: four extinct species known only from fossil remains (three from western North America, one from China), one extant species, the American paddlefish (''Polyodon spathula'') which is native to the
Mississippi River The Mississippi River is the List of longest rivers of the United States (by main stem), second-longest river and chief river of the second-largest Drainage system (geomorphology), drainage system in North America, second only to the Hudson B ...
basin in the U.S., and the Chinese paddlefish (''Psephurus gladius''), declared extinct in 2022 following a 2019 recommendation that it be declared extinct. The species was last sighted in 2003 in the Yangtze River Basin in China. Chinese paddlefish are also commonly referred to as "Chinese swordfish", or "elephant fish". The earliest known species 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 145& ...
( Aptian) 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 stock), resulting in t ...
, 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 past the embryonic stage. Paddlefish have very few bones 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 eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evolut ...
, 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 ancient Greek, βένθος (bénthos), meaning " ...
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 animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, 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 fish Freshwater fish are those that spend some or all of their lives in fresh water, such as rivers and lakes, with a salinity of less than 1.05%. These environments differ from marine conditions in many ways, especially the difference in levels o ...
es in North America, their recorded lengths and weights fall 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 in ''Iowa Fish and Fishing'' (1969) said 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 Kansas () is a U.S. state, state in the Midwestern United States, Midwestern United States. Its Capital city, capital is Topeka, Kansas, Topeka, and its largest city is Wichita, Kansas, Wichita. Kansas is a landlocked state bordered by Nebras ...
on May 5, 2004. However, the record would be broken an additional two times in 2020. On June 28, 2020, an Oklahoma man caught a 146-pounder in Keystone Lake, west of Tulsa. Later on July 23, 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 that paddlefish rostrums are covered in
electroreceptors 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 to stu ...
called
ampullae An ampulla (; ) was, in Ancient Rome, a small round vessel, usually made of glass and with two handles, used for sacred purposes. The word is used of these in archaeology, and of later flasks, often handle-less and much flatter, for holy water or ...
. 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 animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
which is the primary diet of the American paddlefish, but they can also detect the individual feeding and swimming movements of zooplankton's appendages. 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). Therefore, paddlefish with damaged or abbreviated rostrums are still able to forage and maintain good health.


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 in the mid-east region of North America that connect to the Atlantic Ocean via the Saint Lawrence River. There are five lakes ...
and Canada, New York,
Maryland Maryland ( ) is a state in the Mid-Atlantic region of the United States. It shares borders with Virginia, West Virginia, and the District of Columbia to its south and west; Pennsylvania to its north; and Delaware and the Atlantic Ocean t ...
and
Pennsylvania Pennsylvania (; ( Pennsylvania Dutch: )), officially the Commonwealth of Pennsylvania, is a state spanning the Mid-Atlantic, Northeastern, Appalachian, and Great Lakes regions of the United States. It borders Delaware to its southeast, ...
. 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 an arm of the Western Pacific Ocean, located directly offshore from East China. It covers an area of roughly . The sea’s northern extension between mainland China and the Korean Peninsula is the Yellow Sea, separated ...
. 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 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, Red River tributaries, Sabine River, San Jacinto River, Sulphur River, and Trinity River. Their historical range also included occurrences in Canada in Lake Huron and Lake Helen, and in 26–27 states in the United States. The
Ontario Ministry of Natural Resources The Northern Development, Mines, Natural Resources and Forestry is a government ministry of the Canadian province of Ontario that is responsible for Ontario's provincial parks, forests, fisheries, wildlife, mineral aggregates and the Crown lands ...
listed the paddlefish as extirpated from
Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central Ca ...
, Canada 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 the world's most comprehensive inventory of the global conservation status of biolo ...
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, or
CITES CITES (shorter name 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 intern ...
.


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 provided the proper combination of events occur, including water flow, temperature, photoperiod, and availability of gravel substrates suitable for spawning. If all the conditions are not met, paddlefish will 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 spawner Spawn is the eggs and sperm released or deposited into water by aquatic animals. As a verb, ''to spawn'' refers to the process of releasing the eggs and sperm, and the act of both sexes is called spawning. Most aquatic animals, except for aquati ...
s, also referred to as mass spawners or synchronous spawners.
Gravid In biology and human medicine, gravidity and parity are the number of times a woman is or has been pregnant (gravidity) and carried the pregnancies to a viable gestational age (parity). These terms are usually coupled, sometimes with additional t ...
females release their eggs into the water over bare rocks or gravel at the same time males release their sperm. Fertilization occurs externally. 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 US Fish & Wildlife Service and its former
USSR The Soviet Union,. officially the Union of Soviet Socialist Republics. (USSR),. was a transcontinental country that spanned much of Eurasia from 1922 to 1991. A flagship communist state, it was nominally a federal union of fifteen nati ...
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. lot ...
, beginning with five-thousand hatched larvae from
Missouri Missouri is a state in the Midwestern region of the United States. Ranking 21st in land area, it is bordered by eight states (tied for the most with Tennessee): Iowa to the north, Illinois, Kentucky and Tennessee to the east, Arkansas t ...
hatcheries in the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...
. They were introduced into several rivers in Europe and Asia, and provided the first broodstock 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 is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwee ...
,
Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
, the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
, and the Plovdiv and Vidin regions in
Bulgaria Bulgaria (; bg, България, Bǎlgariya), officially the Republic of Bulgaria,, ) is a country in Southeast Europe. It is situated on the eastern flank of the Balkans, and is bordered by Romania to the north, Serbia and North Macedo ...
. Reproduction was successful in 1988 and 1989, and resulted in the exportation of juvenile paddlefish to
Romania Romania ( ; ro, România ) is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern, and Southeast Europe, Southeastern Europe. It borders Bulgaria to the south, Ukraine to the north, Hungary to the west, S ...
and
Hungary Hungary ( hu, Magyarország ) is a landlocked country in Central Europe. Spanning of the Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Cr ...
. In May 2006, specimens of different sizes and weights were caught by professional fisherman near Prahovo in the
Serbia Serbia (, ; Serbian: , , ), officially the Republic of Serbia ( Serbian: , , ), is a landlocked country in Southeastern and Central Europe, situated at the crossroads of the Pannonian Basin and the Balkans. It shares land borders with Hu ...
n part of the
Danube River The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , pa ...
. 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 genus in this family, one recently extinct and three
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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 preserved ...
s: Classification after Grande and Bemis (1991): * Genus †'' Protopsephurus'' Lu, 1994 (Early Cretaceous, China) ** Species †'' Protopsephurus liui'' Lu, 1994 * Genus †'' Paleopsephurus'' MacAlpin, 1947 (Late Cretaceous, North America) ** Species †'' Paleopsephurus wilsoni'' MacAlpin, 1947 * Subfamily Polyodontinae ** Genus †''
Psephurus The Chinese paddlefish (''Psephurus gladius''; : literal translation: "white sturgeon"), also known as the Chinese swordfish, is an extinct species of fish that was formerly native to the Yangtze and Yellow River basins in China. With records of ...
'' Günther, 1873 ***†''
Psephurus gladius The Chinese paddlefish (''Psephurus gladius''; : literal translation: "white sturgeon"), also known as the Chinese swordfish, is an extinct species of fish that was formerly native to the Yangtze and Yellow River basins in China. With records of ...
'' E. von Martens, 1862 Chinese paddlefish (extinct c. 2003) **Tribe Polyodontini ***Genus †'' Crossopholis'' Cope'','' 1883 (
Paleogene The Paleogene ( ; also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period million years ago ( Mya) to the beginning o ...
, North America) ****Species †'' Crossopholis magnicaudatus'' Cope, 1883 ***Genus '' Polyodon'' Lacépède, 1797 (
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pala ...
-Recent, North America) ****'' Polyodon spathula'' Walbaum, 1792 American paddlefish **** †' Grande & Bemis, 1991 Relationships of the genera, after Grande et al. (2002).


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
{{Taxonbar, from=Q858011 Polyodontidae Acipenseriformes Freshwater fish of North America Taxa named by Charles Lucien Bonaparte Extant Barremian first appearances