Family Polyodontidae (Paddlefishes)

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Paddlefishes
Temporal range: Barremian–Recent [1]
American paddlefish, Polyodon spathula
Chinese paddlefish, Psephurus gladius
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Acipenseriformes
Suborder: Acipenseroidei
Family: Polyodontidae
Bonaparte, 1838
Genera

Recent genera

Fossil genera

Paddlefish (family Polyodontidae) are a family of ray-finned fish belonging to the order Acipenseriformes, one of two living groups within this order alongside sturgeons (Acipenseridae).[3][4] Their most distinctive feature is an elongated rostrum that enhances electroreception, allowing them to detect prey in murky water. Often called "primitive fish," paddlefish represent one of the earliest diverging lineages of ray-finned fish, having separated from all other living groups over 300 million years ago. Both living and fossil paddlefish occur almost exclusively in North America and China.[5]

Eight species are known, six of which survive only as fossils—five from North America and one from China.[2] Of the two extant species, the American paddlefish (Polyodon spathula) inhabits the Mississippi River basin in the United States, while the Chinese paddlefish (Psephurus gladius) was declared extinct in 2022 following a 2019 recommendation.[6][7][8] The last confirmed sighting of a Chinese paddlefish occurred in the Yangtze River Basin in 2003.[9][10] The Chinese paddlefish is also known as the "Chinese swordfish" or "elephant fish."[11] The earliest known paddlefish fossil, Protopsephurus, dates to approximately 120 million years ago during the early Cretaceous (Aptian) period in China.

Throughout their historic range, paddlefish populations have declined dramatically due to overfishing, pollution, and human development. Dam construction has proven particularly destructive, blocking seasonal migrations to ancestral spawning grounds. River alterations have changed natural flow patterns, destroying spawning habitat and nursery areas critical to population survival.[12][13]

Morphology

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General morphology of paddlefish

Paddlefish are among the few organisms that retain a notochord beyond the embryonic stage. Their bodies contain very few bones, consisting primarily of cartilage with the notochord serving as a flexible spine. During early development from embryo to fry, paddlefish lack a rostrum (snout), which begins forming shortly after hatching.[14]

The rostrum shape varies dramatically between species. The Chinese paddlefish possessed a narrow, sword-like rostrum, while the American paddlefish displays a broad, paddle-shaped one. Both species share several morphological features: a spindle-shaped body with smooth, scaleless skin, a heterocercal tail, and small, poorly developed eyes.[11][14]

Feeding strategies

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The two species employed vastly different feeding strategies. Unlike its filter-feeding American relative, the Chinese paddlefish was a piscivore—a highly predatory fish. Its forward-pointing jaws suggest it hunted primarily for small fish in the water column, supplementing its diet with shrimp, benthic fishes, and crabs.[11][15] In contrast, the American paddlefish jaw is specialized exclusively for filter feeding. As ram suspension filter feeders, they consume primarily zooplankton, occasionally taking small insects, insect larvae, and small fish.[5]

Size

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Closeup of the head, showing the presence of electrorecepting organs (ampullae of Lorenzini)

The largest recorded Chinese paddlefish measured 23 ft (7.0 m) in length and was estimated to weigh several thousand pounds.[10] More commonly, these fish reached 9.8 ft (3.0 m) and 1,100 lb (500 kg).[10][11][16]

Though the American paddlefish ranks among the largest freshwater fishes in North America, it falls short of its Chinese cousin's impressive dimensions. American paddlefish commonly exceed 5 ft (1.5 m) in length and 60 lb (27 kg) in weight. The largest specimen on record was speared in 1916 in Okoboji Lake, Iowa.[a][17] This fish measured 7 ft 1 in (2.16 m) long with a girth of 45.5 in (1.16 m).[17] According to a 1969 report by J.R. Harlan and E.B. Speaker in Iowa Fish and Fishing, it weighed over 198 lb (90 kg).[18]

The world record paddlefish caught on rod and reel weighed 144 lb (65 kg) and measured 54.25 in (1.378 m) long. Clinton Boldridge caught this fish in a 5-acre pond in Atchison County, Kansas on May 5, 2004.[19][20] This record was broken twice in 2020. On June 28, an Oklahoma man caught a 146-pound paddlefish in Keystone Lake, west of Tulsa. Less than a month later on July 23, another Oklahoma angler caught a 151-pound, nearly 6-foot-long paddlefish in the same lake.[21]

Rostrum and electroreception

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Scientists initially believed paddlefish used their rostrums to dig through bottom sediments,[14][22] but electron microscopy revealed a different purpose entirely. The rostrum surface is covered with electroreceptors called ampullae.[23] These ampullae concentrate densely within star-shaped bone projections that branch from the rostrum.[24]

These electroreceptors detect weak electrical fields that signal prey presence in the water column. Remarkably, they can sense not just zooplankton—the primary food source for American paddlefish—but the individual feeding and swimming movements of zooplankton appendages.[5][14]

Given their poorly developed eyes, paddlefish rely heavily on electroreception for foraging. However, the rostrum is not their only sensory tool. Some sources incorrectly suggest that rostrum damage would severely impair a paddlefish's ability to feed and maintain health. Laboratory experiments and field research demonstrate otherwise. Sensory pores cover nearly half the skin surface, extending from the rostrum across the top of the head down to the tips of the operculum (gill flaps). Even paddlefish with damaged or shortened rostrums can forage effectively.[5][14]

Distribution and habitat

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American paddlefish

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American paddlefish are native to the Mississippi River basin, ranging from New York to Montana and south to the Gulf of Mexico.[25] They inhabit several Gulf Slope drainages, favoring medium to large rivers with long, deep pools that move slowly, along with backwater lakes and bayous.[26] In Texas, paddlefish historically occurred in the Angelina River, Big Cypress Bayou, Neches River, tributaries of the Red River, Sabine River, San Jacinto River, Sulphur River, and Trinity River.[25]

Their historical range once extended into Canada—specifically Lake Huron and Lake Helen—and across 26 to 27 U.S. states. American paddlefish have been extirpated from much of their northern peripheral range, including the Great Lakes and populations in New York, Maryland, Pennsylvania, and Canada. The Ontario Ministry of Natural Resources now lists paddlefish as extirpated from Ontario, Canada under their Endangered Species Act.[27] The IUCN Red List designates Canadian populations as extirpated, noting the last Canadian records date to the early 1900s and the species' presence there was marginal.

Overall, the American paddlefish is classified as vulnerable (VU) on the IUCN Red List. 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).[28] Concerns about remaining populations continue to grow across other states.

Chinese paddlefish

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The Chinese paddlefish was considered anadromous, migrating upstream to spawn, though little is known about their migration patterns or population structure. They were endemic to the Yangtze River Basin in China, living primarily in the broad main stem rivers and shoal zones along the East China Sea.[10][29] Research suggests they favored the middle and lower layers of the water column and occasionally entered large lakes.[10]

With no confirmed sightings since 2003, the species was declared extinct in 2019.[7] Past attempts at artificial propagation for restoration failed because captive specimens could not be kept alive.[30]

Life cycle

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Paddlefish are long-lived and reach sexual maturity late in life. Females do not begin spawning until they reach six to twelve years of age, some not until sixteen to eighteen years old. Males begin spawning around four to seven years of age, with some waiting until nine or ten years.[14][31][4]

Spawning occurs in late spring only when specific environmental conditions align: appropriate water flow, temperature, photoperiod, and the presence of suitable gravel substrates. If any condition is absent, paddlefish will not spawn that year. Females spawn every second or third year, while males spawn more frequently—typically annually or every other year.[14]

To spawn, paddlefish migrate upstream, seeking silt-free gravel bars. During normal conditions, these bars would be exposed to air or covered by very shallow water, but spring snowmelt and rainfall raise river levels sufficiently to flood them.[32] Paddlefish are broadcast spawners, also known as mass or synchronous spawners. Fertilization occurs externally: Gravid females release their eggs over bare rocks or gravel while males simultaneously release sperm. The adhesive eggs stick to the rocky substrate. After hatching, young paddlefish are swept downstream, growing to adulthood in deep freshwater pools.[33]

Aquaculture

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Advances in biotechnology have significantly improved paddlefish propagation and captive rearing. Success rates for reproduction, adaptation, and survival of cultured paddlefish have increased substantially, benefiting both broodstock development and stock rehabilitation programs. These improvements have enabled successful reservoir ranching and pond rearing, generating growing interest in the global market for paddlefish polyculture.[34][35]

In the early 1970s, a cooperative scientific effort between the U.S. Fish & Wildlife Service and its USSR counterpart brought American paddlefish to the former Soviet Union for aquaculture. The program began with five thousand hatched larvae from Missouri hatcheries. These fish were introduced into several European and Asian rivers, establishing the first brood stock that successfully reproduced in 1984–1986 in Russia.[36] Paddlefish are now raised in Germany, Austria, the Czech Republic, and the Plovdiv and Vidin regions in Bulgaria. Reproduction was successful in 1988 and 1989, resulting in the exportation of juvenile paddlefish to Romania and Hungary. In May 2006, specimens of various sizes and weights were caught by professional fishermen near Prahovo in the Serbian part of the Danube River.[36]

In 1988, fertilized paddlefish eggs and larvae from Missouri hatcheries were first introduced into China.[36] Since that time, China imports approximately 4.5 million fertilized eggs and larvae annually from hatcheries in Russia and the United States. Some 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.[35]

Classification

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Restoration of the Cretaceous Protopsephurus
Restoration of Crossopholis

There is one currently extant genus in this family, one recently extinct genus, and five extinct genera known exclusively from fossils.

Classification following Grande & Bemis (1991),[24] with Parapsephurus and Pugiopsephurus added in Hilton et al. (2023):[2]


Relationships of the genera are from Grande et al. (2002).[37]

Polyodontidae

Notes

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  1. ^ "Okoboji Lake" could refer to either of two attached lakes: West Okoboji Lake and East Okoboji Lake. The source does not state which one is meant.

References

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  1. ^ Froese, Rainer; Pauly, Daniel (eds.). "Family Polyodontidae". FishBase. January 2009 version.
  2. ^ a b c d e f g h Hilton, E.J.; During, M.A.D.; Grande, L.; Ahlberg, P.E. (2023). "New paddlefishes (Acipenseriformes, Polyodontidae) from the late Cretaceous Tanis site of the Hell Creek formation in North Dakota, USA". Journal of Paleontology. 97 (3): 675–692. doi:10.1017/jpa.2023.19. S2CID 258095684.
  3. ^ Crow, K.D.; Smith, C.D.; Cheng, J.-F.; Wagner, G.P.; Amemiya, C.T. (2012). "An independent genome duplication inferred from hox paralogs in the American Paddlefish – a representative basal ray-finned fish and important comparative reference". Genome Biology and Evolution. 4 (9): 937–953. doi:10.1093/gbe/evs067. PMC 3509897. PMID 22851613.
  4. ^ a b "Paddlefish (Polyodon spathula)". tpwd.texas.gov. Retrieved 2 December 2022.
  5. ^ a b c d Wilkens, Lon A.; Hofmann, Michael H. (2007). "The paddlefish rostrum as an electrosensory organ: A novel adaptation for plankton feeding". BioScience. 57 (5): 399–407. doi:10.1641/B570505.
  6. ^ "Chinese paddlefish and wild Yangtze sturgeon extinct - IUCN". Reuters. 22 July 2022. Archived from the original on 23 July 2022. Retrieved 22 July 2022.
  7. ^ a b Zhang, Hui; Jarić, Ivan; Roberts, David L.; He, Yongfeng; Du, Hao; Wu, Jinming; et al. (2020). "Extinction of one of the world's largest freshwater fishes: Lessons for conserving the endangered Yangtze fauna". Science of the Total Environment. 710 136242. Bibcode:2020ScTEn.71036242Z. doi:10.1016/j.scitotenv.2019.136242. ISSN 0048-9697. PMID 31911255. S2CID 210086307.
  8. ^ "Study declares ancient Chinese paddlefish extinct". Oceanographic magazine. 9 January 2020. Retrieved 23 April 2022.
  9. ^ "Chinese paddlefish, native to the Yangtze River, declared extinct by scientists". South China Morning Post. 4 January 2020. Retrieved 4 January 2020.
  10. ^ a b c d e Qiwei, W. (2010). "Psephurus gladius". IUCN Red List of Threatened Species. 2010 e.T18428A8264989. doi:10.2305/IUCN.UK.2010-1.RLTS.T18428A8264989.en.
  11. ^ a b c d "Psephurus gladius (Martens, 1862)". Species Fact Sheet. Fisheries and Aquaculture Department. Food and Agriculture Organization of the United Nations. Archived from the original on 7 July 2015. Retrieved 10 June 2014.
  12. ^ "Hooking the dinosaur of fish". The New York Times. 26 May 2018. ISSN 0362-4331. Retrieved 27 May 2018.
  13. ^ "Chinese paddlefish". National Geographic. Archived from the original on 14 July 2014. Retrieved 28 May 2014.
  14. ^ a b c d e f g Biology of the Paddlefish (PDF) (Report). NFC Section I. Lamer-Louisiana State University. Archived from the original (PDF) on 14 July 2014. Retrieved 9 June 2014.
  15. ^ Miller, Michael J. (20 January 2006). "Chapter 4 – The ecology and functional morphology of feeding of North American sturgeon and paddlefish". In le Breton, G.T.O; Beamish, F. William H.; McKinley, Scott R. (eds.). Sturgeons and Paddlefish of North America. Fish & Fisheries Series. Vol. 27. Springer Science & Business Media. pp. 87–101. ISBN 978-1-4020-2833-5. Archived from the original on 28 July 2014. Retrieved 10 June 2014.
  16. ^ Bourton, Jody (29 September 2009). "Giant fish 'verges on extinction'". Earth News. BBC News. Archived from the original on 6 October 2014. Retrieved 9 June 2014.
  17. ^ a b Nichols, J.T. (24 August 1916). "A large Polyodon from Iowa". Copeia. 34 (34): 65. JSTOR 1436920.
  18. ^ Gengerke, Thomas W. (August 1986). "The paddlefish: Status, management and propagation". Distribution and Abundance of Paddlefish in the United States. AFS-7.
  19. ^ "State Record Fish". Kansas angler online edition. Kansas Department of Wildlife, Parks and Tourism. May 2004. Archived from the original on 26 March 2015. Retrieved 9 June 2014.
  20. ^ "State record fish". Kansas Wildlife Parks & Tourism. Retrieved 8 September 2017 – via ksoutdoors.com.
  21. ^ Wilkinson, Joseph (31 July 2020). "Oklahoma man catches world-record 150 pound paddlefish — breaking record set last month in the same lake". The New York Daily News. Retrieved 3 August 2020 – via NYDailyNews.com.
  22. ^ Nachtrieb, Henry F. (1910). "The primitive pores of Polyodon spathula (Walbaum)". Journal of Experimental Zoology. 9 (2): 455–468. doi:10.1002/jez.1400090211 – via zendo.org.
  23. ^ Jørgensen, J. Mørup; Flock, Å.; Wersäll, J. (September 1972). "The Lorenzinian ampullae of Polyodon spathula". Zeitschrift für Zellforschung und Mikroskopische Anatomie. 130 (3): 362–377. doi:10.1007/BF00306949. PMID 4560320. S2CID 28712903.
  24. ^ a b Grande, Lance; Bemis, William E. (28 March 1991). "Osteology and phylogenetic relationships of fossil and recent paddlefishes (Polyodontidae) with comments on the interrelationships of Acipenseriformes". Journal of Vertebrate Paleontology. 11 (sup001): 1–121. doi:10.1080/02724634.1991.10011424. ISSN 0272-4634.
  25. ^ a b "Paddlefish (Polyodon spathula)". Texas Parks & Wildlife. Archived from the original on 5 May 2016. Retrieved 20 April 2016.
  26. ^ INHS padfish (Report). Prairie Research Institute. Urbana-Champaign, IL: University of Illinois. Spring 2002. Archived from the original on 29 May 2014. Retrieved 28 May 2014.
  27. ^ "SAR Paddlefish". Archived from the original on 14 July 2014. Retrieved 9 June 2014.
  28. ^ Grady, J.; et al. (United States Fish and Wildlife Service) (2019). "Polyodon spathula". IUCN Red List of Threatened Species. 2019 e.T17938A174780447. doi:10.2305/IUCN.UK.2019-3.RLTS.T17938A174780447.en. Retrieved 11 November 2021.
  29. ^ "FAO Fisheries & Aquaculture". Retrieved 24 April 2022.
  30. ^ Helfman, Gene (2007). Fish Conservation: A guide to understanding and restoring global aquatic biodiversity and fishery resources. Island Press.
  31. ^ "Paddlefish questions and answers". North Dakota Game and Fish Department. Archived from the original on 29 November 2016. Retrieved 9 June 2014.
  32. ^ "Paddlefish". MDC online. Archived from the original on 11 June 2014. Retrieved 9 June 2014.
  33. ^ Wiley, Edward G. (1998). Paxton, J.R.; Eschmeyer, W.N. (eds.). Encyclopedia of Fishes. San Diego, CA: Academic Press. pp. 77–78. ISBN 0-12-547665-5.
  34. ^ Mims, Steven (2013). Current global status of American paddlefish aquaculture. World Aquaculture Society (meeting abstract). Archived from the original on 3 March 2016. Retrieved 28 April 2015.
  35. ^ a b Mims, Steven D. (February 2006). Paddlefish culture: Development expanding beyond U.S., Russia, China (PDF) (Report). Global Aquaculture Alliance. Archived from the original (PDF) on 14 July 2014. Retrieved 28 April 2015.
  36. ^ a b c Lenhardt, Mirjana; Hegediš, A.; Mićković, B.; Jeftić, Željka Višnjić; Smederevac, Marija; Jarić, I.; et al. (2006). "First record of the North American paddlefish in the Serbian part of the Danube River" (PDF). Arch. Biol. Sci. 58 (3): 27–28. Archived (PDF) from the original on 14 July 2014. Retrieved 9 June 2014 – via doiserbia.nb.rs.
  37. ^ Grande, Lance; Jin, Fan; Yabumoto, Yoshitaka; Bemis, William E. (8 July 2002). "Protopsephurus liui, a well-preserved primitive paddlefish (Acipenseriformes: Polyodontidae) from the lower Cretaceous of China". Journal of Vertebrate Paleontology. 22 (2): 209–237. doi:10.1671/0272-4634(2002)022[0209:plawpp]2.0.co;2. ISSN 0272-4634. S2CID 86258128.
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