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53-595: Lipke Bijdeley Holthuis (21 April 1921 – 7 March 2008) was a Dutch carcinologist , considered one of the "undisputed greats" of carcinology, and "the greatest carcinologist of our time". Holthuis was born in Probolinggo , East Java and obtained his doctorate from Leiden University on 23 January 1946. He was appointed the assistant curator of the Rijksmuseum van Natuurlijke Historie (now Naturalis ) in Leiden in 1941. He

106-417: A Dutch scientist is a stub . You can help Misplaced Pages by expanding it . This article about a zoologist is a stub . You can help Misplaced Pages by expanding it . Crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are a part of the subphylum Crustacea ( / k r ə ˈ s t eɪ ʃ ə / ),

159-403: A large, diverse group of mainly aquatic arthropods including decapods ( shrimps , prawns , crabs , lobsters and crayfish ), 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

212-468: A larger Pancrustacea clade . The traditional classification of Crustacea based on morphology recognised four to six classes. Bowman and Abele (1982) recognised 652 extant families and 38 orders, organised into six classes: Branchiopoda , Remipedia , Cephalocarida , Maxillopoda, Ostracoda , and Malacostraca . Martin and Davis (2001) updated this classification, retaining the six classes but including 849 extant families in 42 orders. Despite outlining

265-468: A limbless abdomen, except from a telson and caudal rami which is present in many groups. The abdomen in malacostracans bears pleopods , and ends in a telson, which bears the anus , and is often flanked by uropods to form a tail fan . The number and variety of appendages in different crustaceans may be partly responsible for the group's success. Crustacean appendages are typically biramous , meaning they are divided into two parts; this includes

318-487: A spiral format. Structures that function as kidneys are located near the antennae. A brain exists in the form of ganglia close to the antennae, and a collection of major ganglia is found below the gut. In many decapods , the first (and sometimes the second) pair of pleopods are specialised in the male for sperm transfer. Many terrestrial crustaceans (such as the Christmas Island red crab ) mate seasonally and return to

371-438: Is an open circulatory system , where blood is pumped into the haemocoel by a heart located near the dorsum. Malacostraca have haemocyanin as the oxygen-carrying pigment, while copepods, ostracods, barnacles and branchiopods have haemoglobins . The alimentary canal consists of a straight tube that often has a gizzard-like "gastric mill" for grinding food and a pair of digestive glands that absorb food; this structure goes in

424-426: Is not too thick it may be possible to see new structures, such as setae , from the outside. However, contact between the nerves and the old exoskeleton is maintained until a very late stage in the process. The new, teneral exoskeleton has to accommodate a larger frame than the previous instar , while the spider has had to fit into the previous exoskeleton until it has been shed. This means the spider does not fill out

477-451: Is produced in Asia, with China alone producing nearly half the world's total. Non-decapod crustaceans are not widely consumed, with only 118,000 tons of krill being caught, despite krill having one of the greatest biomasses on the planet. Ecdysis After moulting, an arthropod is described as teneral , a callow ; it is "fresh", pale and soft-bodied. Within one or two hours,

530-669: Is some debate as to whether or not Cambrian animals assigned to Ostracoda are truly ostracods , which would otherwise start in the Ordovician . The only classes to appear later are the Cephalocarida , which have no fossil record, and the Remipedia , which were first described from the fossil Tesnusocaris goldichi , but do not appear until the Carboniferous . Most of the early crustaceans are rare, but fossil crustaceans become abundant from

583-512: Is then secreted into the exuvial space between the old cuticle and the epidermis, this contains inactive enzymes which are activated only after the new epicuticle is secreted. This prevents the new procuticle from getting digested as it is laid down. The lower regions of the old cuticle, the endocuticle and mesocuticle , are then digested by the enzymes and subsequently absorbed. The exocuticle and epicuticle resist digestion and are hence shed at ecdysis. Spiders generally change their skin for

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636-521: Is thought to be just 1 ⁄ 10 to 1 ⁄ 100 of the total number as most species remain as yet undiscovered . Although most crustaceans are small, their morphology varies greatly and includes both the largest arthropod in the world – the Japanese spider crab with a leg span of 3.7 metres (12 ft) – and the smallest, the 100- micrometre -long (0.004 in) Stygotantulus stocki . Despite their diversity of form, crustaceans are united by

689-611: The Marmorkrebs crayfish. In many crustaceans, the fertilised eggs are released into the water column , while others have developed a number of mechanisms for holding on to the eggs until they are ready to hatch. Most decapods carry the eggs attached to the pleopods , while peracarids , notostracans , anostracans , and many isopods form a brood pouch from the carapace and thoracic limbs. Female Branchiura do not carry eggs in external ovisacs but attach them in rows to rocks and other objects. Most leptostracans and krill carry

742-698: The Carboniferous , as are the first true mantis shrimp. In the Decapoda , prawns and polychelids appear in the Triassic, and shrimp and crabs appear in the Jurassic . The fossil burrow Ophiomorpha is attributed to ghost shrimps, whereas the fossil burrow Camborygma is attributed to crayfishes. The Permian–Triassic deposits of Nurra preserve the oldest (Permian: Roadian) fluvial burrows ascribed to ghost shrimps (Decapoda: Axiidea, Gebiidea) and crayfishes (Decapoda: Astacidea, Parastacidea), respectively. However,

795-465: The Carboniferous period onwards. Within the Malacostraca, no fossils are known for krill , while both Hoplocarida and Phyllopoda contain important groups that are now extinct as well as extant members (Hoplocarida: mantis shrimp are extant, while Aeschronectida are extinct; Phyllopoda: Canadaspidida are extinct, while Leptostraca are extant ). Cumacea and Isopoda are both known from

848-401: The Japanese spider crab with a leg span of up to 3.8 m (12.5 ft) and a mass of 20 kg (44 lb). Like other arthropods , crustaceans have an exoskeleton , which they moult to grow. They are distinguished from other groups of arthropods, such as insects , myriapods and chelicerates , by the possession of biramous (two-parted) limbs, and by their larval forms , such as

901-503: The hexapods ( insects and entognathans ) emerged deep in the Crustacean group, with the completed pan-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 ( oligostracans and multicrustaceans ). The 67,000 described species range in size from Stygotantulus stocki at 0.1 mm (0.004 in), to

954-629: The nauplius stage of branchiopods and copepods . Most crustaceans are free-living aquatic animals , but some are terrestrial (e.g. woodlice , sandhoppers ), some are parasitic (e.g. Rhizocephala , fish lice , tongue worms ) and some are sessile (e.g. barnacles ). The group has an extensive fossil record , reaching back to the Cambrian . More than 7.9 million tons of crustaceans per year are harvested by fishery or farming for human consumption, consisting mostly of shrimp and prawns . Krill and copepods are not as widely fished, but may be

1007-413: The pereon or thorax , and the pleon or abdomen . The head and thorax may be fused together to form a cephalothorax , which may be covered by a single large carapace . The crustacean body is protected by the hard exoskeleton , which must be moulted for the animal to grow. The shell around each somite can be divided into a dorsal tergum , ventral sternum and a lateral pleuron. Various parts of

1060-405: The prosoma with sufficient pressure to crack it open along its lines of weakness. The carapace lifts off from the front, like a helmet, as its surrounding skin ruptures, but it remains attached at the back. Now the spider works its limbs free and typically winds up dangling by a new thread of silk attached to its own exuviae, which in turn hang from the original silk attachment. At this point

1113-515: The zoea (pl. zoeæ or zoeas ). This name was given to it when naturalists believed it to be a separate species. It follows the nauplius stage and precedes the post-larva . Zoea larvae swim with their thoracic appendages , as opposed to nauplii, which use cephalic appendages, and megalopa, which use abdominal appendages for swimming. It often has spikes on its carapace , which may assist these small organisms in maintaining directional swimming. In many decapods , due to their accelerated development,

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1166-702: The Asian shore crab, Hemigrapsus sanguineus . Since the opening of the Suez Canal , close to 100 species of crustaceans from the Red Sea and the Indo-Pacific realm have established themselves in the eastern Mediterranean sub-basin, with often significant impact on local ecosystems. Most crustaceans have separate sexes , and reproduce sexually . In fact, a recent study explains how the male T. californicus decide which females to mate with by dietary differences, preferring when

1219-683: The Hexapoda is distinctly closer to e.g. a Multicrustacean than an Oligostracan is. Crustaceans have a rich and extensive fossil record , which begins with animals such as Canadaspis and Perspicaris from the Middle Cambrian age Burgess Shale . Most of the major groups of crustaceans appear in the fossil record before the end of the Cambrian, namely the Branchiopoda , Maxillopoda (including barnacles and tongue worms ) and Malacostraca ; there

1272-406: The animals with the greatest biomass on the planet, and form a vital part of the food chain. The scientific study of crustaceans is known as carcinology (alternatively, malacostracology , crustaceology or crustalogy ), and a scientist who works in carcinology is a carcinologist . The body of a crustacean is composed of segments, which are grouped into three regions: the cephalon or head ,

1325-436: The back allowing the animal to emerge. Often, this initial crack is caused by a combination of movement and increase in pressure of hemolymph within the body, forcing an expansion across its exoskeleton , leading to an eventual crack that allows for certain organisms such as spiders to extricate themselves. While the old cuticle is being digested, the new layer is secreted. All cuticular structures are shed at ecdysis, including

1378-420: The cuticle and collapse of air sacs to allow growth of internal organs. The process of moulting in insects begins with the separation of the cuticle from the underlying epidermal cells (apolysis) and ends with the shedding of the old cuticle (ecdysis). In many species it is initiated by an increase in the hormone ecdysone . This hormone causes: After apolysis the insect is known as a pharate . Moulting fluid

1431-643: The cuticle hardens and darkens following a tanning process analogous to the production of leather . During this short phase the animal expands, since growth is otherwise constrained by the rigidity of the exoskeleton. Growth of the limbs and other parts normally covered by the hard exoskeleton is achieved by transfer of body fluids from soft parts before the new skin hardens. A spider with a small abdomen may be undernourished but more probably has recently undergone ecdysis. Some arthropods, especially large insects with tracheal respiration, expand their new exoskeleton by swallowing or otherwise taking in air. The maturation of

1484-580: The description of 428 new taxa: 2 new families, 5 subfamilies, 83 genera and 338 species. 67 taxa were named after him between 1953 ( Hippolyte holthuisi ) and 2009 ( Caridina holthuisi , Lysmata holthuisi ). However, in Fransen, C.H.J.M., De Grave, S., Ng, P.K.L. 2010, an additional 50 taxa were named after him. In 1972 Holthuis received an honorary doctorate from the Norwegian University of Science and Technology (NTNU) . This article about

1537-448: The eggs between their thoracic limbs; some copepods carry their eggs in special thin-walled sacs, while others have them attached together in long, tangled strings. Crustaceans exhibit a number of larval forms, of which the earliest and most characteristic is the nauplius . This has three pairs of appendages , all emerging from the young animal's head, and a single naupliar eye. In most groups, there are further larval stages, including

1590-679: The evidence that Maxillopoda was non-monophyletic, they retained it as one of the six classes, although did suggest that Maxillipoda could be replaced by elevating its subclasses to classes. Since then phylogenetic studies have confirmed the polyphyly of Maxillipoda and the paraphyletic nature of Crustacea with respect to Hexapoda. Recent classifications recognise ten to twelve classes in Crustacea or Pancrustacea, with several former maxillopod subclasses now recognised as classes (e.g. Thecostraca , Tantulocarida , Mystacocarida , Copepoda , Branchiura and Pentastomida ). The following cladogram shows

1643-444: The exoskeleton may be fused together. Each somite , or body segment can bear a pair of appendages : on the segments of the head, these include two pairs of antennae , the mandibles and maxillae ; the thoracic segments bear legs , which may be specialised as pereiopods (walking legs) and maxillipeds (feeding legs). Malacostraca and Remipedia (and the hexapods) have abdominal appendages. All other classes of crustaceans have

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1696-456: The females are algae-fed instead of yeast-fed. A small number are hermaphrodites , including barnacles , remipedes , and Cephalocarida . Some may even change sex during the course of their life. Parthenogenesis is also widespread among crustaceans, where viable eggs are produced by a female without needing fertilisation by a male. This occurs in many branchiopods , some ostracods , some isopods , and certain "higher" crustaceans, such as

1749-454: The females before maturing. Members of the Mygalomorphae are very long-lived, sometimes 20 years or more; they moult annually even after they mature. Spiders stop feeding at some time before moulting, usually for several days. The physiological processes of releasing the old exoskeleton from the tissues beneath typically cause various colour changes, such as darkening. If the old exoskeleton

1802-410: The first time while still inside the egg sac, and the spiderling that emerges broadly resembles the adult. The number of moults varies, both between species and sexes, but generally will be between five times and nine times before the spider reaches maturity. Not surprisingly, since males are generally smaller than females, the males of many species mature faster and do not undergo ecdysis as many times as

1855-662: The great radiation of crustaceans occurred in the Cretaceous , particularly in crabs, and may have been driven by the adaptive radiation of their main predators, bony fish . The first true lobsters also appear in the Cretaceous. Many crustaceans are consumed by humans, and nearly 10,700,000  tons were harvested in 2007; the vast majority of this output is of decapod crustaceans : crabs , lobsters , shrimp , crawfish , and prawns . Over 60% by weight of all crustaceans caught for consumption are shrimp and prawns, and nearly 80%

1908-557: The inner parts of the exoskeleton, which includes terminal linings of the alimentary tract and of the tracheae if they are present. Each stage of development between moults for insects in the taxon Endopterygota is called an instar , or stadium, and each stage between moults of insects in the Exopterygota is called a nymph: there may be up to 15 nymphal stages. Endopterygota tend to have only four or five instars. Endopterygotes have more alternatives to moulting, such as expansion of

1961-422: The layer allow light to reach the retina. As the larvae mature into adults, the layer migrates to a new position behind the retina where it works as a backscattering mirror that increases the intensity of light passing through the eyes, as seen in many nocturnal animals. In an effort to understand whether DNA repair processes can protect crustaceans against DNA damage , basic research was conducted to elucidate

2014-426: The limb is a normal, or near normal, size. The term ecdysis comes from Ancient Greek ἐκδύω ( ekduo ) 'to take off, strip off'. In preparation for ecdysis, the arthropod becomes inactive for a period of time, undergoing apolysis or separation of the old exoskeleton from the underlying epidermal cells. For most organisms, the resting period is a stage of preparation during which the secretion of fluid from

2067-417: The moulting glands of the epidermal layer and the loosening of the underpart of the cuticle occurs. Once the old cuticle has separated from the epidermis, a digesting fluid is secreted into the space between them. However, this fluid remains inactive until the upper part of the new cuticle has been formed. Then, by crawling movements, the organism pushes forward in the old integumentary shell , which splits down

2120-488: The new exoskeleton completely, so it commonly appears somewhat wrinkled. Most species of spiders hang from silk during the entire process, either dangling from a drop line, or fastening their claws into webbed fibres attached to a suitable base. The discarded, dried exoskeleton typically remains hanging where it was abandoned once the spider has left. To open the old exoskeleton, the spider generally contracts its abdomen ( opisthosoma ) to supply enough fluid to pump into

2173-401: The oceans as insects are on land. Most crustaceans are also motile , moving about independently, although a few taxonomic units are parasitic and live attached to their hosts (including sea lice , fish lice , whale lice , tongue worms , and Cymothoa exigua , all of which may be referred to as "crustacean lice"), and adult barnacles live a sessile life – they are attached headfirst to

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2226-429: The repair mechanisms used by Penaeus monodon (black tiger shrimp). Repair of DNA double-strand breaks was found to be predominantly carried out by accurate homologous recombinational repair. Another, less accurate process, microhomology-mediated end joining , is also used to repair such breaks. The expression pattern of DNA repair related and DNA damage response genes in the intertidal copepod Tigriopus japonicus

2279-446: The sea to release the eggs. Others, such as woodlice , lay their eggs on land, albeit in damp conditions. In most decapods, the females retain the eggs until they hatch into free-swimming larvae. Most crustaceans are aquatic, living in either marine or freshwater environments, but a few groups have adapted to life on land, such as terrestrial crabs , terrestrial hermit crabs , and woodlice . Marine crustaceans are as ubiquitous in

2332-510: The second pair of antennae, but not the first, which is usually uniramous , the exception being in the Class Malacostraca where the antennules may be generally biramous or even triramous. It is unclear whether the biramous condition is a derived state which evolved in crustaceans, or whether the second branch of the limb has been lost in all other groups. Trilobites , for instance, also possessed biramous appendages. The main body cavity

2385-473: The special larval form known as the nauplius . The exact relationships of the Crustacea to other taxa are not completely settled as of April 2012 . Studies based on morphology led to the Pancrustacea hypothesis, in which Crustacea and Hexapoda ( insects and allies) are sister groups . More recent studies using DNA sequences suggest that Crustacea is paraphyletic , with the hexapods nested within

2438-501: The spider is a callow; it is teneral and vulnerable. As it dangles, its exoskeleton hardens and takes shape. The process may take minutes in small spiders, or some hours in the larger Mygalomorphs. Some spiders, such as some Synema species, members of the Thomisidae (crab spiders), mate while the female is still callow, during which time she is unable to eat the male. Eurypterids are a group of chelicerates that became extinct in

2491-403: The structure and colouration of the new exoskeleton might take days or weeks in a long-lived insect; this can make it difficult to identify an individual if it has recently undergone ecdysis. Ecdysis allows damaged tissue and missing limbs to be regenerated or substantially re-formed. Complete regeneration may require a series of moults, the stump becoming a little larger with each moult until

2544-500: The substrate and cannot move independently. Some branchiurans are able to withstand rapid changes of salinity and will also switch hosts from marine to non-marine species. Krill are the bottom layer and most important part of the food chain in Antarctic animal communities. Some crustaceans are significant invasive species , such as the Chinese mitten crab, Eriocheir sinensis , and

2597-412: The updated relationships between the different extant groups of the paraphyletic Crustacea in relation to the class Hexapoda . Ostracoda Mystacocarida Branchiura Pentastomida Malacostraca Copepoda Tantulocarida Thecostraca Cephalocarida Branchiopoda   Remipedia Hexapoda According to this diagram, the Hexapoda are deep in the Crustacea tree, and any of

2650-426: The zoea is the first larval stage. In some cases, the zoea stage is followed by the mysis stage, and in others, by the megalopa stage, depending on the crustacean group involved. Providing camouflage against predators, the otherwise black eyes in several forms of swimming larvae are covered by a thin layer of crystalline isoxanthopterin that gives their eyes the same color as the surrounding water, while tiny holes in

2703-447: Was analyzed after ultraviolet irradiation. This study revealed increased expression of proteins associated with the DNA repair processes of non-homologous end joining , homologous recombination , base excision repair and DNA mismatch repair . The name "crustacean" dates from the earliest works to describe the animals, including those of Pierre Belon and Guillaume Rondelet , but the name

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2756-412: Was not used by some later authors, including Carl Linnaeus , who included crustaceans among the " Aptera " in his Systema Naturae . The earliest nomenclatural valid work to use the name "Crustacea" was Morten Thrane Brünnich 's Zoologiæ Fundamenta in 1772, although he also included chelicerates in the group. The subphylum Crustacea comprises almost 67,000 described species , which

2809-486: Was the most prolific carcinologist of the 20th century, publishing 620 papers (108 of which were in the Leiden Museum Journals ) totalling 12,795 pages which is an average of 185 pages per year and an average of approximately 21 pages per paper. These were published on many groups of crustaceans , their natural history and nomenclature, and the history of carcinology. This steady stream of publications resulted in

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