Triops longicaudatus

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Triops longicaudatus
Triops-longicaudatus-dorsal-ventral-edit2.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Branchiopoda
Subclass: Phyllopoda
Superorder: Calmanostraca
Order: Notostraca
Family: Triopsidae
Genus: Triops
Species:
T. longicaudatus
Binomial name
Triops longicaudatus
(LeConte, 1846)
Triopsrange.PNG
The range of Triops longicaudatus
Synonyms

Apus longicaudatusLeConte, 1846

Triops longicaudatus (commonly called American tadpole shrimp or longtail tadpole shrimp) 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.

Contents

Classification

Triops longicaudatus is a member of the crustacean class Branchiopoda, which primarily contains freshwater animals with gills on their legs. The class Branchiopoda is divided into the subclasses Sarsostraca, containing fairy shrimp, and Phyllopoda, containing all other members (cladocerans, clam shrimps, and the tadpole shrimp). The subclass name, literally meaning leaf-footed, is derived from their flat, leaflike appendages. Notostracans are placed in the infraclass Calamanostraca, which also contains the extinct Kazacharthra. [1]

Structure

Comparison of T. longicaudatus with T. cancriformis
Triops longicaudatus.jpg
Triops longicaudatus
Triops cancriformis2.jpg
Triops cancriformis

Triops longicaudatus is usually greyish yellow or brown in colour, and differs from many other species by the absence of the second maxilla. Apart from Triops cancriformis , it is the only tadpole shrimp species whose individuals display as many as three reproductive strategies: bisexual, unisexual (parthenogenetic), and hermaphroditic; see below. Triops cancriformis is easily recognizable by its yellow carapace with dark spots, whilst T. longicaudatus individuals have a uniform carapace. As its name suggests, its elongated tail structures (cercopods) are often nearly as long as the rest of the body; including the cercopods, the body may reach 75 mm (3.0 in) in length. [2]

External anatomy of Triops longicaudatus (click on image for captions) Triops anatomy new.png
External anatomy of Triops longicaudatus (click on image for captions)

The head of Triops longicaudatus is typical of crustaceans and consists of five segments, but there is a tendency toward reduction of cephalic appendages. The trunk is not distinctly divided into thorax and abdomen. Most trunk segments bear appendages. Zoologists find it difficult to decide where the thorax stops and the abdomen begins; the debate is seemingly endless. The first eleven trunk segments each bear a single pair of limbs. They are followed by segments that are fused similarly to those of millipedes, and as a result, each segment bears up to six pairs of limbs. The trunk ends with a region of limbless segments. Some zoologists consider the thorax to consist of the two regions with appendages, and the abdomen the region without appendages. Others believe the region of fused segments to be part of the abdomen. [3] For clarity, this article uses "thorax" to mean the two body regions with limbs.

Carapace

There is a carapace present, but no cephalothorax, since no thoracic segment is fused with the head. The terms carapace and cephalothorax often are confused, but should not be. The carapace of crustaceans is a fold of the body wall of the fifth head segment.

Mouthparts: 1 - ventral groove; 2 - labrum (upper lip); 3 - mandible Mouthparts.jpg
Mouthparts: 1 – ventral groove; 2 – labrum (upper lip); 3 – mandible

The head bears a pair of dorsal compound eyes that lie close to each other and are nearly fused together. The compound eyes are generally sessile (not stalked). In addition, there is a naupliar ocellus (the "third eye") between them. The compound eyes are on the surface of the head, but the ocellus is deep within the head. All the eyes, however, are easily visible through the shell covering of the head. A distinct horizontal groove, known as the mandibular groove, marks the division between the anterior three head segments and the posterior two. Posterior to it is the cervical groove, marking the division between the head and the thorax.

On the ventral side of the head is a lenslike window, admitting light to the naupliar eye, which is aimed both dorsally and ventrally. The first antennae (antennules) are small, slender filaments on the ventral surface of the head, at about the same level as the eyes. The second antennae are similar and lie laterally to the first. They are nonfunctioning. The large, well-developed mandibles oppose each other across the ventral midline. The opposing surfaces bear strong brownish teeth. As the crustacean periodically opens and closes the mandibles the teeth move apart and close together. Of the usual crustacean head appendages, only the mandibles are well developed. In Triops longicaudatus, the larger second maxillae are absent, only maxillules being present. [3]

Thorax

Magnified leg (thoracopod): 1-5 - endites; 6 - endopod; 7 - exopod; 8 - epipod; 9 - protopod A thoracopod.jpg
Magnified leg (thoracopod): 1–5 – endites; 6 – endopod; 7 – exopod; 8 – epipod; 9 – protopod

The anterior thorax consists of eleven segments, each bearing a pair of appendages, called thoracopods or pereiopods. None of the thoracopods are modified into maxillipeds for feeding. The posterior thorax consists of 16–25 segments, incompletely separated to form rings. Each ring may consist of as many as six fused segments and bear up to six pairs of appendages (there are 54–66 limbs altogether).

Most of the thoracic appendages resemble each other, but the first eleven pairs are best developed. There is a slight tendency to regional specialization and the first pair of legs is unlike the rest. It is an elongated, cheliped-like structure with a sensory function, while all the other appendages aid in feeding, respiration, and locomotion. In females, the eleventh pair of legs is modified into brood pouches. The many legs posterior to the eleventh pair move the spent feeding and respiratory currents away. [3]

Abdomen

The posterior 5–14 rings at the end of the body do not bear appendages. At the end of the body is a pleotelson (fusion of the last abdominal segment and the telson) connected to long, multisegmented uropods. [3]

Feeding behaviour

The feeding method of tadpole shrimps is similar to that proposed for the ancestral crustacean. The anterior appendages (second pair to tenth pair) stir sediments and swirl muddy water into the wide, midventral food groove. The gnathobases (inward-facing lobes at the base of the leg) guide food anteriorly to the mouth. The large flat exopods (outward-facing lobes at the end of the leg) stir and lift the sediments. Fine silt particles and water escape easily, but large, coarse food particles are torn into smaller pieces by the blade-like, inward-facing lobes called endopods at the end of the leg.

Internal anatomy

The heart of tadpole shrimps is a long dorsal tube in the anterior eleven trunk segments. It has a pair of ostia in each of these segments. Sometimes hemoglobin is present in its blood and the crustacean may be pink as a result. The excretory organs are the paired maxillary glands, located on the segment of the second maxilla. The long looped ducts of these glands can be seen in the carapace. The role of the maxillary glands is primarily osmoregulatory. Nitrogen, in the form of ammonia, is lost by diffusion across the gill surfaces. [3]

Reproduction and development

Reproduction

Vernal Pools 2.jpg
A typical vernal pool during the wet season
Vernal Pools 3.jpg
Another vernal pool, this one dried out

Triopslongicaudatus displays several reproductive strategies. Individuals may reproduce sexually, but this is rare, as most populations are highly male- or female-biased. Parthenogenesis (development from unfertilized eggs) is the most common reproductive strategy. Some populations, however, consist of hermaphrodites who fertilize each other. Different populations display different strategies or combinations of strategies, and therefore, in the future may be considered separate species or subspecies.

In females, the eleventh pair of legs is modified into egg sacs, where the eggs are carried for several hours. The eggs are released in batches, have a thick shell, and can stand freezing temperatures as well as drought, enabling the population to survive from one season to the next. The eggs have to dry out completely before being submerged in water again in order to hatch successfully; they may remain in a state of diapause for up to 20 years. These eggs may have helped Triops longicaudatus, as well as other notostracans, to survive the various natural disasters and mass extinctions to date. [1] [4]

Life cycle

To complete their lives, tadpole shrimps depend on the changing nature of the temporary waters they inhabit. During the dry season (summer and autumn), their offspring stay inside the eggs. As the pool fills with rainwater during the winter and spring, they hatch and feed on fairy shrimps and other invertebrates. The first larval stage (the metanauplius) is orange in colour. It has a single eye, six legs, and develops through instars (growth stages). Each instar ends with shedding the exoskeleton. The number of segments and appendages increases as Triops grow, and they slowly change to greyish brown. In approximately eight days, they reach maturity and lay eggs. Adult Triops die as the pools dry up. Triops generally live for about 20–90 days if the pool does not dry up. [5]

Larval development (not to scale)
Tadpole shrimp cyst hatching.jpg
Metanauplius hatches from the egg, having three pairs of limbs: the antennules, antennae and mandibles.
Tadpole shrimp larva 2.jpg
Three hours later, the trunk segments become defined and the telson begins to form.
Tadpole shrimp larva 1.jpg
7 h after hatching
Tadpole shrimp larva 3.jpg
After 15 h, it has the major characteristics of the adult form.
Tadpole shrimp larva 4.jpg
24 h after hatching, it very nearly resembles a miniature version of the adult form.
Triops Longicaudatus at about 48 hours old.jpg
Two days after hatching, the Triops has essentially completely taken the appearance of an adult.

Ecology

Red variant of Triops longicaudatus raised in captivity Triopslongred1.jpg
Red variant of Triops longicaudatus raised in captivity

T. longicaudatus is the most widespread notostracan species, and may be found in western North America, South America, Japan, South Korea, and several Pacific Islands. It is most active at a temperature of approximately 20 °C (68 °F), and is usually found scratching the mud at the bottom of pools, searching for benthic food. Triops collect food particles by straining the water with hairs on their limbs. Loose food particles are collected in a groove running down the underside of the body lengthwise, and held together by a sticky secretion until they are swallowed by their very small (2 mm wide) mouth. The tiny mouth is deep in its underbelly, and while the animal is capable of breaking up plant roots or dead fish, it is incapable of chasing down and eating prey larger than it is.

Diet

A Triops eating its companion Triops longicaudatus eating his companion - Cannibalism.jpg
A Triops eating its companion

Tadpole shrimps are omnivorous and may eat algae, insects, and other organic debris; they are known to chase very small fry, tadpoles, and oligochaete worms. [3] In general, they eat anything organic that is smaller than they are, which even may include their siblings (they are cannibalistic). In turn, Triops longicaudatus are eaten by frogs and birds. [6]

Interaction with humans

Adult 2.5 cm pet Triops in aquarium Triops longicaudatus2.jpg
Adult 2.5 cm pet Triops in aquarium

The species is considered a human ally against the West Nile virus, as the individuals consume Culex mosquito larvae. They also are used as a biological agent in Japan, eating weeds in rice paddies. In Wyoming, the presence of Triops longicaudatus usually indicates a good chance of the hatching of spadefoot frogs.

Dried eggs of Triopslongicaudatus are sold in kits to be raised as aquarium pets, sold under the name of "aquasaurs", "trigons" or "triops". These are most often of the greenish-brown variety, but the red variant is fairly common among enthusiasts. Captive Triops are frequently kept in aquariums and fed a diet consisting mainly of carrots, shrimp pellets and dried shrimp. Often they are also given living shrimp and daphnia as live prey. Because they can feed on just about anything, they are also fed lunch meat, potatoes, crackers, etc. They will often feed themselves by grazing on algae and other grunge from the bottom and sides of the tank and on small particles clinging to sponge filters or to Marimo moss balls, which are often cultured alongside Triops.

Distribution

Though they make fine captive tank mates, Triops longicaudatus red and Triops cancriformis are not likely to be found together in the wild, nor to feed on carrots. Marimo moss balls are also unlikely companions. Triopslongredcanc.jpg
Though they make fine captive tank mates, Triops longicaudatus red and Triops cancriformis are not likely to be found together in the wild, nor to feed on carrots. Marimo moss balls are also unlikely companions.

Triops longicaudatus is widespread in North America. In Canada, it is found only in the provinces of Alberta, Saskatchewan and Manitoba. [7] It is widespread throughout the contiguous United States, Mexico, and Hawaii, but not Alaska. Tadpole shrimps may be found in parts of South America (Argentina and Galápagos Islands), [8] in the West Indies, and the Pacific Islands, including Japan and New Caledonia. [9]

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 and the Devonian Lepidocaris. 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 crustaceans 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">Mysida</span> Small, shrimp-like crustacean

Mysida is an order of small, shrimp-like crustaceans in the malacostracan superorder Peracarida. Their common name opossum shrimps stems from the presence of a brood pouch or "marsupium" in females. The fact that the larvae are reared in this pouch and are not free-swimming characterises the order. The mysid's head bears a pair of stalked eyes and two pairs of antennae. The thorax consists of eight segments each bearing branching limbs, the whole concealed beneath a protective carapace and the abdomen has six segments and usually further small limbs.

<span class="mw-page-title-main">Isopoda</span> Order of arthropods

Isopoda is an order of crustacean, which includes woodlice and their relatives. Members of this group are called Isopods and include both terrestrial and aquatic species. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, and five pairs of branching appendages on the abdomen that are used in respiration. Females brood their young in a pouch under their thorax.

<span class="mw-page-title-main">Notostraca</span> Order of small freshwater animals

The order Notostraca, containing the single family Triopsidae, is a group of crustaceans known as tadpole shrimp or shield shrimp. 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.

<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">Cephalocarida</span> Class of crustaceans

The Cephalocarida are a class in the subphylum Crustacea comprising only 12 species. Both the nauplii and the adults are benthic. They were discovered in 1955 by Howard L. Sanders, and are commonly referred to as horseshoe shrimp. They have been grouped together with the Remipedia in the Xenocarida. Although a second family, Lightiellidae, is sometimes used, all cephalocaridans are generally considered to belong in just one family: Hutchinsoniellidae. Fossil records of cephalocaridans has been found in 462 million year old deposits.

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<i>Canadaspis</i> Extinct genus of arthropods

Canadaspis is an extinct genus of bivalved Cambrian arthropod, known from North America and China. They are thought to have been benthic feeders that moved mainly by walking and possibly used its biramous appendages to stir mud in search of food. They have been placed within the Hymenocarina, which includes other bivalved Cambrian arthropods.

Eucrenonaspides oinotheke is a species of crustacean in the family Psammaspidae, endemic to Tasmania, the only species described in the genus Eucrenonaspides. The Eucrenonaspides is a variation of the Anaspidaceans. It was described from a spring at 9 Payton Place, Devonport, Tasmania in 1980, making it "the first spring-dwelling syncarid recorded from the Australian region". It is listed as a vulnerable species on the IUCN Red List. A further undescribed species is known from south-western Tasmania.

<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>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 belong to the subphylum Crustacea, and form a large, diverse group of 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.

References

  1. 1 2 Grzimek's Animal Life Encyclopedia. Volume 02 – Protostomes. Chapter 14 – Order Notostraca
  2. June Latting & Peter Glanville Rowlands, ed. (1995). The California Desert: an Introduction to Natural Resources and Man's Impact. Vol. 2. Riverside, CA: June Latting. p. 278. ISBN   9780963690852. OCLC   37199208.
  3. 1 2 3 4 5 6 Richard Fox (June 19, 2006). "Triops longicaudatus, Tadpole Shrimp". Invertebrate Anatomy OnLine. Lander University. Archived from the original on October 1, 2010. Retrieved October 4, 2010.
  4. David Wooten. "Triops longicaudatus". Zooplankton of the Great Lakes. Central Michigan University . Retrieved October 4, 2010.
  5. Encarta Encyclopedia 2005. Article – Branchiopoda
  6. Allison Batdorff (June 29, 2005). "Wet weather revives crustaceans prehistoric heritage". Billings Gazette.
  7. Dale Parker. "Saskatchewan Macrocrustacea" . Retrieved October 4, 2010.
  8. "Class Branchiopoda" (PDF). biota.org.br/. Archived from the original (PDF) on 2012-01-04.
  9. "Species Descriptions". MyTriops.com. Archived from the original on October 6, 2011. Retrieved October 4, 2010.