Eunicidae ist der Name einer Familie kleiner bis sehr großer meist räuberischer, selten parasitischer Vielborster (Polychaeta), die in Meeren weltweit entweder frei lebend, röhrenbildend oder grabend zu finden sind.
Die Vielborster der Familie Eunicidae haben bis zu 1500 Segmente und werden zwischen 1 cm und 6 m lang. Der segmentierte Körper ist nicht in größere Abschnitte gegliedert, jedoch verändern sich Größe und Gestalt der Parapodien an den Seiten der Segmente entlang des Körpers. Kennzeichnend für die Familie sind die am zweilappigen oder ungeteilten Prostomium (Kopflappen) befindlichen 1 bis 3 Antennen, die keine geringelten Antennenträger (Ceratophoren) besitzen. In der Gattung Eunice sind es 2 Palpen und 3 Antennen, also 5 Fortsätze am Prostomium, während es in anderen Gattungen weniger sein können. Die Palpen (buccalen Lippen) können reduziert oder auch wohl entwickelt sein. Die meisten Arten der Eunicidae besitzen Augen. Das Peristomium besteht aus zwei Ringen, an deren hinterem bei manchen Arten zwei peristomiale Cirren sitzen. Die Parapodien sind verzweigt und das Notopodium nur als dorsaler Cirrus ausgebildet. Die Kiemen, soweit vorhanden, bestehen aus einfachen oder kammartigen Filamenten. Die Neuropodien haben obere deutlich gekantete und kammartige Borsten. Die Kiefer bestehen aus ventralen unverschmolzenen Mandibeln und dorsalen, mit Aragonit mineralisierten Maxillen, die aus einem Paar kurzer Träger und 4 bis 5 Platten rechts sowie 5 bis 6 Platten links zusammengesetzt sind.
Der Körperbau ist bei den Arten dieser Familie sehr einheitlich, so dass insbesondere Form und Anzahl der Fortsätze am Prostomium sowie Anzahl und Verteilung der Kiemen für die Bestimmung wichtig sind. Wie die meisten großen Polychaeten besitzen die Eunicidae in ihrem geschlossenen Blutgefäßsystem zur Bindung des Sauerstoffs als Blutfarbstoff Hämoglobine, die frei im Blut gelöst und nicht an Blutkörperchen gebunden sind. Auf Grund ihrer durch das Blut rot gefärbten parapodialen Kiemen werden viele Arten auch als „Blutwürmer“ bezeichnet.
Weibchen und Männchen sind bei den Eunicidae gleich groß. Die meisten Arten haben in ihren Segmenten Mixonephridien mit einem einfachen Ausscheidungskanal, der gleichzeitig als Ausgang für die Gameten dient. Die Eizellen sind mindestens 0,1 mm groß und bieten genug Lecithin für eine Larve, mehrere Tage ohne zu fressen als Plankton zu leben. Soweit bekannt, dauert das Larvenstadium etwa 3 bis 8 Tage bis zur Metamorphose zum kriechenden Wurm. Direkte Entwicklung ohne Larvenstadium ist in der Familie unbekannt. Bei den meisten Arten leben die Jungwürmer nach der Metamorphose zunächst frei kriechend in Felsspalten, ehe sie bei vielen Arten zu einem Leben als Röhrenwurm übergehen.
Eine der größten und am besten untersuchten Arten der Eunicidae ist die sowohl im Indischen Ozean als auch im Pazifischen Ozean unter anderem an der Küste Südaustraliens verbreitete bis zu 2 m lange Eunice aphroditois, die in langen Röhren im Substrat lebt und nach vorbeischwimmenden Tieren schnappt, wobei sie neben einer Vielzahl anderer Beutetiere auch Fische frisst. Die ebenfalls im Pazifik, insbesondere um Samoa, die Fidschi-Inseln sowie die Kleinen Sundainseln verbreitete, ebenso sehr große Palola viridis ernährt sich dagegen von den in Korallen lebenden symbiontischen Algen. Ähnlich wie andere Vertreter der weltweit in warmen Gewässern verbreiteten Gattung Palola ist ihr Fortpflanzungsverhalten an die Mondphasen gebunden. Der als Epitoke bezeichnete hintere Abschnitt des Körpers, der die Gonaden enthält, wird zur Fortpflanzungszeit abgeschnürt und bewegt sich an die Wasseroberfläche, wo er aufplatzt und so die Gameten ins freie Meerwasser entlässt. Durch die Vielzahl an nebeneinander treibenden männlichen und weiblichen Epitoken ist die äußere Befruchtung im Meerwasser sichergestellt. Es entwickeln sich frei schwimmende Trochophora-Larven.
Trotz ihrer teilweise großen Wehrhaftigkeit bilden Borstenwürmer der Familie Eunicidae die Hauptnahrung verschiedener räuberisch lebender Schneckenarten, darunter die Stachelschnecke Drupa morum,[1] die Keramikvase (Vasum ceramicum)[2] und die Schwarzweiße Kegelschnecke (Conus ebraeus).[3]
Die Familie Eunicidae wird in elf Gattungen unterteilt:[4]
Eunicidae ist der Name einer Familie kleiner bis sehr großer meist räuberischer, selten parasitischer Vielborster (Polychaeta), die in Meeren weltweit entweder frei lebend, röhrenbildend oder grabend zu finden sind.
Eunicidae is a family of marine polychaetes (bristle worms). The family comprises marine annelids distributed in diverse benthic habitats across Oceania, Europe, South America, North America, Asia and Africa.[1] The Eunicid anatomy typically consists of a pair of appendages near the mouth (mandibles) and complex sets of muscular structures on the head (maxillae) in an eversible pharynx.[2] One of the most conspicuous of the eunicids is the giant, dark-purple, iridescent "Bobbit worm" (Eunice aphroditois), a bristle worm found at low tide under boulders on southern Australian shores. Its robust, muscular body can be as long as 2 m.[3] Eunicidae jaws are known from as far back as Ordovician sediments.[4][5] Cultural tradition surrounds Palola worm (Palola viridis) reproductive cycles in the South Pacific Islands.[6] Eunicidae are economically valuable as bait in both recreational and commercial fishing.[7][8] Commercial bait-farming of Eunicidae can have adverse ecological impacts.[9] Bait-farming can deplete worm and associated fauna population numbers,[10] damage local intertidal environments [11] and introduce alien species to local aquatic ecosystems.[12]
In 2020, Zanol et al. stated, "Species traditionally considered to belong to Eunice are now, also, distributed in two other genera Leodice and Nicidion recently resurrected to reconcile Eunicidae taxonomy with its phylogenetic hypothesis."[13]
In 1992, Kristian Fauchald detailed a conclusive history of research and classification of the Eunicidae family.[4] Primary studies undertaken in 1767 on coral reefs in Norway, initially classified Eunicid species under the Nereis family.[4] In 1817, Georges Cuvier created a new genus, Eunice, to classify these and other original taxa.[4] Throughout the 1800s (1832-1878) worm species were added to this genera by Jean Victor Audouin and Henri Milne-Edwards, Kinberg, Edwardsia de Quatrefages, Malmgren, Ehlers and Grube.[4] Following the Challenger and Albatross expeditions, research was expanded by McIntosh and Chamberlain.[4] In 1921 and 1922, Treadwell added new species from coral reefs in the Caribbean Sea and the Pacific Ocean.[4] Species were reviewed and their classifications were refined by Fauvel, Augener and Hartman throughout the early 1900s.[4] In 1944, Hartman codified a system of separate classification for the family, informally grouping North American species using the original suggestions of Ehlers.[4] Hartman's system was expanded and specified by Fauchald in 1970 and later again by Miura in 1986.[4]
Thirty-three genera have been described in the Eunicidae family.[14] Only eight are currently considered valid:[15]
Eunicids can contain up to 1500 segments and span from 10 mm to 6 m in length. Members of the Eunicidae family are distinguished from other families in Eunicida by having a rear segment with 1-3 antennae and no ringed bases on their antennae.[16] The first body segment of Eunicidae is either whole or consists of two lobes.[16] The gills of live specimens are typically identifiable by their bright red colour.[17]
A pair of slender and cylindrical sensory appendages are typically situated near the head of Eunicidae.[16] The lips of Eunicidae can be either reduced or well-developed.[16] In the Eunice species, worms have five appendages on two elongated segmented appendages and three antennae near their heads.[16] This feature is not part of the anatomy of all genera in the Eunicidae family. Eunicidae jaws are typically well developed and partly visible on the underside of the worm or on its surface at the front of the mouth in a complex structure.[16][17]
Some species of Eunicidae have extensions of the body wall that loop into the vascular system.[16] These usually consist of either comb-like or single filaments.[16]
Eunicidae are distributed in diverse benthic habitats across Oceania, Europe, South America, North America, Asia and Africa. Eunicids play an ecological role in benthic communities, exhibiting a preference for subtidal hard substrates in shallow temperate waters, tropical waters and mangrove swamps.[1][4] Most species of Eunicidae inhabit cracks and crevices in assorted rubble, rock, and sand environments.[4] In limestone or coral reefs, Eunicids burrow into hard parchment-like tube corals or remain in crevices of calcareous algae.[18]
Eunicid diets vary across genera. For example, the Eunice aphroditois crawl on the seafloor where they scavenge in a carnivorous feeding pattern on marine worms, small crustaceans, molluscs, algae and detritus.[2][14][19][20][21] Other species, for example Euniphysa tubifex and large Eunice, hunt the surrounds of their coral habitats and feed on the decaying flesh of dead sea-life.[2][22] Burrowing species of Eunicidae (Lysidice and Palola) are primarily herbivores. These species feed on matured corals and contained organisms or on types of algae.[23] The diet of Marphysa species of Eunicidae is variable, some worms are herbivores,[21] some are carnivores [24] and others omnivores.[2][22]
The practice of harvesting polychaetes (including species in the Eunicidae family) as bait may have negative ecological impacts on intertidal habitats and on worm population numbers.[9][11] In 2019, Cabral et al. found that Marphysa sanguinea are placed at risk by overfishing and unlicensed harvesting in Portugal.[9] The ecological impacts of bait harvesting activity can also affect associated fauna populations [10] as well as sediment quality [25] and bioavailability of heavy metals.[26][9] Research indicates that mudworm survival and growth may also be affected by changes in salinity rates.[27]
Importing Eunicidae species is an established alternative to exploiting local populations for bait.[12] This process may lead to accidental species introductions or invasions.[28][29] Alien species can threaten the foundation of local ecosystems by altering food webs, habitat structures and gene pools.[28] Alien species can also introduce diseases and parasites.[29][30] Six species of Eunice, one species of Euniphysa, three species of Lysidice and one species of Marphysa sp. were identified as alien in local aquatic ecosystems across the Mediterranean, the Red Sea, the USA Pacific and the North Sea.[28] Live bait worms are often emptied into the water body by anglers at the end of a fishing session, this is another practice that can introduce alien species to aquatic ecosystems.[12][28][29]
Most of the class Polychaeta are benthic sexual reproductive animals and lack external reproductive organs.[31] When mating, female polychaetes produce a pheromone that induces a mutual release of male sperm and female eggs. This process of synchronous reproduction in the form of a swarm is known as epitoky. During this process, there is no actual male to female contact. The reproductive swarm is ejected into open water. Cells that fuse during fertilisation (gametes) are spawned through an excretory gland (metanephridia) or by the main worm body-wall rupturing.[32] Post-fertilisation, most eggs become planktonic; although some remain inside the worm tubes or burrow in external jelly masses attached to the tubes.[32] Epitokes can draw an increased number of pelagic predators.[6] In the Florida Keys for example, the swarming of Eunice fucata is a highly publicised in local fishing communities, attracting a large gathering of tarpon.[6] These mass swarming events, or ‘risings’, are a spectacle that is the foundation of local tradition in Samoa, Fiji, Tonga, Papua New Guinea, Vanuatu, Kiribati and Indonesia.[1]
Marphysa sanguinea, or known locally in Italy as “Murrido”, “Murone”, “Bacone” and “Verme sanguigno” is the most valuable bait of all Polychaete species collected in Italy.[7] This species is also cultivated in USA and South Korea and is typically commercially harvested once at its optimal length of 20–30 cm.[7] Marphysa sanguinea can reach up to 50 cm long and is collected by excavating in deep sediment.[8] For example, in the Venice lagoon, fisherman dig below the sediment layers colonised by the nereidids and sieve organic material through coarse screens.[33] This process is also common in Italian coastal areas with intertidal and shallow littoral muddy bottoms.[7] Eunice aphroditois, another sizeable (up to 1 metre in length) species of Eunicidae, is harvested by scuba divers along the Italian Apulia coasts.[7] This species is collected at soft bottom ocean floors at a depth of 10 metres using specialised harvesting instruments that fit into U-shaped parchment tubes where the worm lives.[8] This species of Eunicidae is suitable bait for fish of the Sparidae family and is used in commercial hook and line practice.[7] Species within the Eunicidae family are also caught by recreational and commercial fisherman in estuaries along the West coast of Portugal and in Arcahon Bay in France.[7][34] Marphysa are propagated and harvested in Australian estuary communities located along the coast of New South Wales and Queensland.[34] Collecting of Marphysa moribidii as bait occurs along the West coast of Peninsular Malaysia, Marphysa elityeni are caught in subsistence fisheries in Africa and Eunice sebastiani have been reported as being harvested for bait in Brazil.[34] Eunicids are also used as supplementary feed for aquaculture.[8][35][36][37] For example, mudworms are a part of the black tiger prawn diet in some Thailand hatcheries.[35][38]
In the Indo-Pacific, during 1-2 nights each year, the epitokes of the Palola viridis species are automatised.[1][6] The sizeable epitokes (up to 30 cm in length) swim autonomously upwards and rupture, releasing gametes across the surface of the ocean.[1] The epitokes are composed of hundreds of segments, with females emerald in colour and males transitioning from orange to brown during maturation.[6] On ‘rising’ night it is tradition for some local communities to attract epitokes with artificial light sources or using other traditional methods.[34] In Samoa for example, locals wear necklaces made of mosoʻoi flowers and use the fragrant floral scent to attract Palola worms.[34] The epitokes are scooped from the shallows into nets and containers to be consumed raw, or cooked, baked, dried or frozen for later consumption.[34]
Eunicidae is a family of marine polychaetes (bristle worms). The family comprises marine annelids distributed in diverse benthic habitats across Oceania, Europe, South America, North America, Asia and Africa. The Eunicid anatomy typically consists of a pair of appendages near the mouth (mandibles) and complex sets of muscular structures on the head (maxillae) in an eversible pharynx. One of the most conspicuous of the eunicids is the giant, dark-purple, iridescent "Bobbit worm" (Eunice aphroditois), a bristle worm found at low tide under boulders on southern Australian shores. Its robust, muscular body can be as long as 2 m. Eunicidae jaws are known from as far back as Ordovician sediments. Cultural tradition surrounds Palola worm (Palola viridis) reproductive cycles in the South Pacific Islands. Eunicidae are economically valuable as bait in both recreational and commercial fishing. Commercial bait-farming of Eunicidae can have adverse ecological impacts. Bait-farming can deplete worm and associated fauna population numbers, damage local intertidal environments and introduce alien species to local aquatic ecosystems.
In 2020, Zanol et al. stated, "Species traditionally considered to belong to Eunice are now, also, distributed in two other genera Leodice and Nicidion recently resurrected to reconcile Eunicidae taxonomy with its phylogenetic hypothesis."
Los eunícidos (Eunicidae) son gusanos marinos poliquetos, muchos de los cuales alcanzan un tamaño considerable. Sus mandíbulas son conocidas en sedimentos desde el Ordovícico.[1]
Son depredadores, algunos se alimentan de coral o son parásitos. Poseen una probóscide eversible. Uno de los más llamativos es el "gusano Bobbit" (Eunice aphroditois), un poliqueto gigante de color morado oscuro, iridiscente, que se observa durante la marea baja bajo las rocas de las costas del sur de Australia; su cuerpo robusto y musculoso puede llegar hasta los 3 m.[2]
Los eunícidos (Eunicidae) son gusanos marinos poliquetos, muchos de los cuales alcanzan un tamaño considerable. Sus mandíbulas son conocidas en sedimentos desde el Ordovícico.
Son depredadores, algunos se alimentan de coral o son parásitos. Poseen una probóscide eversible. Uno de los más llamativos es el "gusano Bobbit" (Eunice aphroditois), un poliqueto gigante de color morado oscuro, iridiscente, que se observa durante la marea baja bajo las rocas de las costas del sur de Australia; su cuerpo robusto y musculoso puede llegar hasta los 3 m.
Les Eunicidae sont une famille de vers annélides polychètes marins. Leurs fossiles sont connus depuis l'Ordovicien[2]. Nombreux sont les eunicides à atteindre une taille considérable (2 mètres).
Selon World Register of Marine Species (24 nov. 2012)[3] :
Selon ITIS (24 nov. 2012)[4] :
Palola sp.
Les Eunicidae sont une famille de vers annélides polychètes marins. Leurs fossiles sont connus depuis l'Ordovicien. Nombreux sont les eunicides à atteindre une taille considérable (2 mètres).
Eunicidae adalah salah satu famili dari Polychaeta. Banyak eunicidae yang dapat mencapai ukuran yang besar. Rahang eunicidae dapat diketahui dari sedimen ordovician[1]. Eunicidae hidup di laut; beberapa dari mereka merupakan parasit.[1]
Salah satu spesies yang paling mencolok dari eunicidae ialah Eunice aphroditois, yang mempunyai ukuran raksasa, berwarna ungu tua, ditemukan di air pasang surut di bawah batu-batu di pesisir Australia. Eunice aphroditois mempunyai badan yang berotot dan cukup kuat, serta dapat mencapai panjang 2 meter[2].
Beberapa spesies eunicidae memangsa karang, dan juga telah ditemukan individu yang hidup di karang akuarium untuk waktu yang cukup lama dan tumbuh raksasa[3][4].
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(bantuan) Eunicidae adalah salah satu famili dari Polychaeta. Banyak eunicidae yang dapat mencapai ukuran yang besar. Rahang eunicidae dapat diketahui dari sedimen ordovician. Eunicidae hidup di laut; beberapa dari mereka merupakan parasit.
Salah satu spesies yang paling mencolok dari eunicidae ialah Eunice aphroditois, yang mempunyai ukuran raksasa, berwarna ungu tua, ditemukan di air pasang surut di bawah batu-batu di pesisir Australia. Eunice aphroditois mempunyai badan yang berotot dan cukup kuat, serta dapat mencapai panjang 2 meter.
Beberapa spesies eunicidae memangsa karang, dan juga telah ditemukan individu yang hidup di karang akuarium untuk waktu yang cukup lama dan tumbuh raksasa.
Gli Eunicidi (Eunicidae Berthold, 1827) sono una famiglia di anellidi policheti.
La famiglia comprende i seguenti generi:
Uno dei più grossi eunicidi è la gigantesca, purpurea, iridescente Eunice aphroditois, che può essere trovata anche sulla spiaggia con la bassa marea lungo le coste meridionali dell'Australia. Il suo corpo robusto e muscoloso può raggiungere 1 m di lunghezza.[1]
Eunicidae zijn een familie van borstelwormen (Polychaeta).
De volgende geslachten zijn bij de familie ingedeeld:
Eunicidae zijn een familie van borstelwormen (Polychaeta).
Латинское название Eunicidae Berthold, 1827[1] ITIS 66260 NCBI 104737
Eunicidae (лат.) — семейство морских многощетинковых червей из отряда Eunicida [2]. Имеют от одной до пяти затылочных антенн[3][4]. Ископаемые остатки известны из Ордовика. Встречаются повсеместно в морях; несколько видов ведут паразитический образ жизни [5]. Один из наиболее крупных видов — пурпурный австралийский Eunice aphroditois — достигает в длину до 2-3 м[6]. Некоторые виды поедают кораллы. Отдельные особи могут жить в рифовых аквариумах, долгое время оставаясь незамеченными[7][8].
Eunicidae (лат.) — семейство морских многощетинковых червей из отряда Eunicida . Имеют от одной до пяти затылочных антенн. Ископаемые остатки известны из Ордовика. Встречаются повсеместно в морях; несколько видов ведут паразитический образ жизни . Один из наиболее крупных видов — пурпурный австралийский Eunice aphroditois — достигает в длину до 2-3 м. Некоторые виды поедают кораллы. Отдельные особи могут жить в рифовых аквариумах, долгое время оставаясь незамеченными.
Aciculomarphysa Hartmann-Schröder & Zibrowius, 1998 Eunice Cuvier, 1817 Euniphysa Wesenberg-Lund, 1949 Fauchaldius Carrera-Parra & Salazar Vallejo, 1998 Lysibranchia Cantone, 1983 Lysidice Lamarck, 1818 Marphysa Quatrefages, 1865 Nematonereis Schmarda, 1861 Palola Gray in Stair, 1847