Notostraca

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Notostraca
Temporal range: Famennian–Recent
Triops australiensis.JPG
Triops australiensis
LepidurusApus.jpg
Lepidurus apus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Branchiopoda
Superorder: Calmanostraca
Order: Notostraca
G. O. Sars, 1867
Genera
  • Lepidurus Leach, 1819
  • Triops Schrank, 1803
  • Fossil genera and species, see text

The order Notostraca, containing the single family Triopsidae, is a group of crustaceans known as tadpole shrimp [1] or shield shrimp. [2] The two genera, Triops and Lepidurus , are considered living fossils, with similar forms having existed since the end of the Devonian, around 360 million years ago. They have a broad, flat carapace, which conceals the head and bears a single pair of compound eyes. The abdomen is long, appears to be segmented and bears numerous pairs of flattened legs. The telson is flanked by a pair of long, thin caudal rami. Phenotypic plasticity within taxa makes species-level identification difficult, and is further compounded by variation in the mode of reproduction. Notostracans are omnivores living on the bottom of temporary pools and shallow lakes.

Contents

Description

Notostracans are 2–5 centimetres (0.8–2.0 in) long, with a broad carapace at the front end, and a long, slender abdomen. [1] This gives them a similar overall shape to a tadpole, from which the common name tadpole shrimp derives. [1] The carapace is dorso-ventrally flattened, smooth, and bears no rostrum; it includes the head, and the two sessile compound eyes are located together on top of the head. [1] The two pairs of antennae are much reduced, with the second pair sometimes missing altogether. [2] The mouthparts comprise a pair of uniramous mandibles and no maxillipeds. [2]

The ventral side of Triops australiensis, showing the many pairs of phyllopodous legs Triops australiensis belly.jpg
The ventral side of Triops australiensis, showing the many pairs of phyllopodous legs

The trunk consists of three regions; thorax I, thorax II and the abdomen. Thorax I is made up of 11 segments, each with a pair of well-developed limbs and the genital opening on the eleventh segment. In the female, it is modified to form a "brood pouch". [3] The first one or two pairs of legs differ from the remainder, and probably function as sensory organs. [3]

The somites on thorax II are fused into "rings", which varies in number between species and gender and appear to be body segments, but do not always reflect the underlying segmentation. [1] Each ring is made up of 2–6 complete or partial fused segments, and the number of legs on each body ring match its number of segments. [4] [1] The legs become progressively smaller posteriorly, [3] with the last segments being legless. [1]

The limbless abdomen ends in a telson and a pair of long, thin, multi-articulate caudal rami. [5] The form of the telson varies between the two genera: in Lepidurus , a rounded projection extends between the caudal rami, while in Triops there is no such projection. [1]

Life cycle

A collection of lake sediment containing the pink eggs of Triops longicaudatus Collected Eggs of Triops Longicaudatus.JPG
A collection of lake sediment containing the pink eggs of Triops longicaudatus

Within the Notostraca, and even within species, there is variation in the mode of reproduction, with some populations reproducing sexually, some showing self-fertilisation of females, and some showing a mix of the two. [1] [6] The frequency of males in populations is therefore highly variable. [3] In sexual populations, the sperm leave the male's body through simple pores, there being no penis. The eggs are released by the female and then held in the cup-like brood pouch. [3] The eggs are retained by the female only for a short time before being laid, [7] and the larvae develop directly, without passing through a metamorphosis. [2]

Ecology and distribution

Notostracans are omnivorous, eating small animals such as fishes and fairy shrimp. [1] They are found worldwide in freshwater, brackish water, or saline pools, as well as in shallow lakes, peat bogs, and moorland. [2] The species Triops longicaudatus is considered an agricultural pest in California rice paddies, because it prevents light from reaching the rice seedlings by stirring up sediment. [8]

Evolution and fossil record

The fossil record of Notostraca is extensive, occurring in a wide range of geological deposits. [9] The oldest known notostracan is the species Strudops goldenbergi from the Late Devonian (Famennian ~ 365 million years ago) of Belgium. [10] The lack of major morphological change since 250  million years ago has led to Notostraca being described as living fossils. [11] Kazacharthra, a group known only from Triassic and Jurassic fossils from Kazakhstan and Western China, [12] are closely related to notostracans, and may belong within the order Notostraca, [13] or alternatively are placed as their sister group within the clade Calmanostraca.

The "central autapomorphy" of the Notostraca is the abandonment of filter feeding in open water, and the development of a benthic lifestyle in muddy waters, taking up food from particles of sediment and preying on small animals. [7] A number of other characteristics are correlated with this change, including the increased size of the animal compared to its relatives, and the loss of the ability to hinge the carapace; although a central keel marks the former separation into two valves, the adductor muscle is missing. [7] Notostracans retain the plesiomorphic condition of having two separate compound eyes, which abut, but have not become united, as seen in other groups of Branchiopoda. [7]

Taxonomy

The extant members of order Notostraca composed a single family, Triopsidae, with only two genera, Triops and Lepidurus . [11] The problematic Middle Ordovician fossil Douglasocaris has been erected and placed in its own family Douglasocaridae by Caster & Brooks 1956, and may be ancestral to Notostraca.

The phenotypic plasticity shown by notostracan species make identification to the species level difficult. [11] Many putative species have been described based on morphological variation, such that by the 1950s, as many as 70 species were recognised. [11] Two important revisions – those of Linder in 1952 [14] and Longhurst in 1955 [15] synonymised many taxa, and resulted in the recognition of only 11 species in the two genera. This taxonomy was accepted for decades, [11] "even attaining the status of dogma". [16] More recent studies, especially those employing molecular phylogenetics, have shown that the eleven currently recognised species conceal a greater number of reproductively isolated populations. [11]

Genera list

Incertae sedis species

See also

Related Research Articles

<span class="mw-page-title-main">Branchiopoda</span> Class of crustaceans

Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca, Notostraca, the Devonian Lepidocaris and possibly the Cambrian Rehbachiella. They are mostly small, freshwater animals that feed on plankton and detritus.

<span class="mw-page-title-main">Malacostraca</span> Largest class of crustaceans

Malacostraca is the second largest of the six classes of pancrustaceans just behind hexapods, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, amphipods, mantis shrimp, tongue-eating lice and many other less familiar animals. They are abundant in all marine environments and have colonised freshwater and terrestrial habitats. They are segmented animals, united by a common body plan comprising 20 body segments, and divided into a head, thorax, and abdomen.

<span class="mw-page-title-main">Clam shrimp</span> Suborder of arthropods

Clam shrimp are a group of bivalved branchiopod crustaceans that resemble the unrelated bivalved molluscs. They are extant and also known from the fossil record, from at least the Devonian period and perhaps before. They were originally classified in the former order Conchostraca, which later proved to be paraphyletic, due to the fact that water fleas are nested within clam shrimps. Clam shrimp are now divided into three orders, Cyclestherida, Laevicaudata, and Spinicaudata, in addition to the fossil family Leaiidae.

<span class="mw-page-title-main">Anostraca</span> Order of crustaceans

Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered mountain lakes, and Antarctic ice. They are usually 6–25 mm (0.24–0.98 in) long. Most species have 20 body segments, bearing 11 pairs of leaf-like phyllopodia, and the body lacks a carapace. They swim "upside-down" and feed by filtering organic particles from the water or by scraping algae from surfaces, with the exception of Branchinecta gigas, or "giant fairy shrimp", which is itself a predator of other species of anostracans. They are an important food for many birds and fish, and some are cultured and harvested for use as fish food. There are 300 species spread across 8 families.

<i>Triops</i> Genus of small crustaceans

Triops is a genus of small crustaceans in the order Notostraca. The long-lasting resting eggs of several species of Triops are commonly sold in kits as pets. The animals hatch upon contact with fresh water. Most adult-stage Triops have a life expectancy of up to 90 days and can tolerate a pH range of 6 to 10. In nature, they often inhabit temporary pools.

<span class="mw-page-title-main">Tanaidacea</span> Order of crustaceans

The crustacean order Tanaidacea make up a minor group within the class Malacostraca. There are about 940 species in this order.

<i>Triops longicaudatus</i> Species of small freshwater animal

Triops longicaudatus is a freshwater crustacean of the order Notostraca, resembling a miniature horseshoe crab. It is characterized by an elongated, segmented body, a flattened shield-like brownish carapace covering two thirds of the thorax, and two long filaments on the abdomen. The genus name Triops comes from Greek ὤψ or ṓps, meaning "eye" prefixed with Latin tri-, "three", in reference to its three eyes. Longicaudatus is an Latin neologism combining longus ("long") and caudatus ("tailed"), referring to its long tail structures. Triops longicaudatus is found in fresh water ponds and pools, often in places where few higher forms of life can exist.

<i>Triops australiensis</i> Species of small freshwater animal

Triops australiensis, sometimes referred to as a shield shrimp, is an Australian species of the tadpole shrimp Triops.

<i>Triops newberryi</i> Species of small freshwater animal

Triops newberryi is a species of Triops found on the western coast of North America, commonly in valleys throughout the states of Washington, Oregon, California, and small areas of Nevada, Utah, New Mexico, and Mexico, with at least one disjunct population in Kansas. They are found in vast numbers though in the Coachella Valley in California. T. newberryi has been reported to have potential as a biocontrol agent for larval mosquitoes breeding in seasonally-flooded habitats. T. newberryi is genetically distinct from T. longicaudatus, the dominant species in the Central United States.

<i>Triops cancriformis</i> Species of small freshwater animal

Triops cancriformis, European tadpole shrimp or tadpole shrimp is a species of tadpole shrimp found in Europe to the Middle East and India.

<span class="mw-page-title-main">Crustacean larva</span> Crustacean larval and immature stages between hatching and adult form

Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The larvae of crustaceans often bear little resemblance to the adult, and there are still cases where it is not known what larvae will grow into what adults. This is especially true of crustaceans which live as benthic adults, more-so than where the larvae are planktonic, and thereby easily caught.

<i>Chenops</i> Genus of small freshwater animals

Chenops is an extinct genus of notostracan which existed in the Yixian Formation, Inner Mongolia, and the Jehol fauna of China during the early Cretaceous period. The genus was erected by Thomas A. Hegna and Ren Dong in 2010 to describe the Yixian species, Chenops yixianensis. A second species, originally described as "Prolepidurus oblongus", from the Jehol fauna, was redescribed as C. oblongus.

<i>Jeholops</i> Genus of small freshwater animals

Jeholops is an extinct genus of notostracan which existed in the Yixian Formation, inner Mongolia, China during the early Cretaceous period. It was described by Thomas A. Hegna and Ren Dong in 2010, and the only species is Jeholops hongi.

<i>Chirocephalus diaphanus</i> Species of small freshwater animal

Chirocephalus diaphanus is a widely distributed European species of fairy shrimp that lives as far north as Great Britain, where it is the only surviving species of fairy shrimp and is protected under the Wildlife and Countryside Act 1981. It is a translucent animal, about 0.5 in (13 mm) long, with reddened tips to the abdomen and appendages. The body comprises a head, a thorax bearing 11 pairs of appendages, and a seven-segmented abdomen. In males, the antennae are enlarged to form "frontal appendages", while females have an egg pouch at the end of the thorax.

<i>Lepidocaris</i> Extinct genus of crustaceans

Lepidocaris rhyniensis is an extinct species of crustacean. It is the only species known from the order Lipostraca, and is the only abundant animal in the Pragian-aged Rhynie chert deposits. It resembles modern Anostraca, to which it is probably closely related, although its relationships to other orders remain unclear. The body is 3 mm (0.12 in) long, with 23 body segments and 19 pairs of appendages, but no carapace. It occurred chiefly among charophytes, probably in alkaline temporary pools.

<span class="mw-page-title-main">Crustacean</span> Subphylum of arthropods

Crustaceans are a group of arthropods that are a part of the subphylum Crustacea, a large, diverse group of mainly aquatic arthropods including decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, opossum shrimps, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea. The three classes Cephalocarida, Branchiopoda and Remipedia are more closely related to the hexapods than they are to any of the other crustaceans.

<i>Lepidurus apus</i> Species of small freshwater animal

Lepidurus apus, commonly known as a tadpole shrimp, is a notostracan in the family Triopsidae, one of a lineage of shrimp-like crustaceans that have had a similar form since the Triassic period and are considered living fossils. This species is cosmopolitan, inhabiting temporary freshwater ponds over much of the world, and the most widespread of the tadpole shrimps. Like other notostracans, L. apus has a broad carapace, long segmented abdomen, and large numbers of paddle-like legs. It reproduces by a mixture of sexual reproduction and self-fertilisation of females.

<i>Tyrannophontes</i> Extinct genus of mantis shrimp

Tyrannophontes is an extinct genus of mantis shrimp that lived during the late Carboniferous period in what is now the Mazon Creek fossil beds of Illinois. It is the only genus in the family Tyrannophontidae. The type species, T. theridion, was described in 1969 by Frederick Schram. A second, much larger species, T. gigantion, was also named by Schram in 2007. Two other species were formerly assigned to the genus, but have since been reclassified.

Nodosculda is an extinct genus of mantis shrimp that lived in North America during the late Albian stage of the Early Cretaceous period, between 105 and 100 million years ago. The only species is Nodosculda fisherorum, known from several specimens uncovered in the Paw Paw Formation of Texas.

References

  1. 1 2 3 4 5 6 7 8 9 10 Denton Belk (2007). "Branchiopoda". In Sol Felty Light; James T. Carlton (eds.). The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon (4th ed.). University of California Press. pp. 414–417. ISBN   978-0-520-23939-5.
  2. 1 2 3 4 5 J. K. Lowry (October 2, 1999). "Notostraca (Branchiopoda)". Crustacea, the Higher Taxa: Description, Identification, and Information Retrieval. Archived from the original on July 23, 2011. Retrieved February 7, 2011.
  3. 1 2 3 4 5 Douglas Grant Smith (2001). Pennak's freshwater invertebrates of the United States: Porifera to Crustacea (4th ed.). John Wiley and Sons. ISBN   978-0-471-35837-4.
  4. Evolution and Phylogeny of Pancrustacea: A Story of Scientific Method - 44. Notostraca and Allies (Calmanostraca)
  5. "Subclass 1. Branchiopoda". The Invertebrata (4th ed.). Cambridge University Press. 1961. pp. 368–375.
  6. Self-Fertilization and the Role of Males in Populations of Tadpole Shrimp (Branchiopoda: Notostraca: Triops)
  7. 1 2 3 4 Peter Ax (2000). "Notostraca". Multicellular Animals. The Phylogenetic System of the Metazoa. Volume II. Springer. pp. 158–159. ISBN   978-3-540-67406-1.
  8. Clifford, Hugh F. (1991). "Notostraca: Tadpole shrimp". Aquatic Invertebrates of Alberta: An illustrated guide. University of Alberta. pp. 144–145. ISBN   978-0-88864-234-9.
  9. Atte Korhola & Milla Rautio (2001). "Cladocera and other branchiopod crustaceans". In John P. Smol; Harry John Betteley Birks & William M. Last (eds.). Tracking Environmental Change Using Lake Sediments. Volume 4: Zoological Indicators. Kluwer Academic Publishers. pp. 5–41. ISBN   978-1-4020-0658-6.
  10. Lagebro, Linda; Gueriau, Pierre; Hegna, Thomas A.; Rabet, Nicolas; Butler, Aodhán D.; Budd, Graham E. (May 2015). Korn, Dieter (ed.). "The oldest notostracan (Upper Devonian Strud locality, Belgium)". Palaeontology. 58 (3): 497–509. Bibcode:2015Palgy..58..497L. doi: 10.1111/pala.12155 . ISSN   0031-0239. S2CID   129231634.
  11. 1 2 3 4 5 6 Luc Brendonck; D. Christopher Rogers; Jorgen Olesen; Stephen Weeks & Walter R. Hoch (2008). "Global diversity of large branchiopods (Crustacea: Branchiopoda) in freshwater". In Estelle V. Balian; Christian Lévêque; Hendrik Segers & Koen Martens (eds.). Freshwater Animal Diversity Assessment. Developments in Hydrobiology 198. Vol. 595. pp. 167–176. doi:10.1007/s10750-007-9119-9. ISBN   978-1-4020-8258-0. S2CID   46608816.{{cite book}}: |journal= ignored (help)
  12. Liu, Hongfu (1996). "New materials of Late Triassic Kazacharthra from Xinjaing". Acta Palaeontologica Sinica. 4 (35): 490–494. Archived from the original on 2013-04-14.
  13. Marjorie L. Reaka-Kudla (2002). "Habitat specialization and its relation to conservation policy in Crustacea". In Elva Escobar-Briones; Fernando Alvarez (eds.). Modern Approaches to the Study of Crustacea. Springer. pp. 211–221. ISBN   978-0-306-47366-1.
  14. Folke Linder (1952). "Contributions to the morphology and taxonomy of the Branchiopoda Notostraca, with special reference to the North American species". Proceedings of the United States National Museum . 102 (3291): 1–69. doi:10.5479/si.00963801.102-3291.1.
  15. Alan R. Longhurst (1955). "A review of the Notostraca". Bulletin of the British Museum (Natural History) . 3 (1): 1–57. doi: 10.5962/bhl.part.4119 .
  16. Clay Sassaman; Marie A. Simovich & Michael Fugate (1997). "Reproductive isolation and genetic differentiation in North American species of Triops (Crustacea: Branchiopoda: Notostraca)". Hydrobiologia . 359 (1–3): 125–147. doi:10.1023/A:1003168416080. S2CID   6767325.
  17. Wagner, Philipp; Haug, Joachim T.; Sell, Jürgen; Haug, Carolin (December 2017). "Ontogenetic sequence comparison of extant and fossil tadpole shrimps: no support for the "living fossil" concept". PalZ. 91 (4): 463–472. Bibcode:2017PalZ...91..463W. doi:10.1007/s12542-017-0370-8. ISSN   0031-0220. S2CID   90922613.
  18. 1 2 Wagner, Philipp; Haug, Joachim T.; Haug, Carolin (December 2019). "A new calmanostracan crustacean species from the Cretaceous Yixian Formation and a simple approach for differentiating fossil tadpole shrimps and their relatives". Zoological Letters. 5 (1): 20. doi: 10.1186/s40851-019-0136-0 . ISSN   2056-306X. PMC   6582493 . PMID   31245037.