Priapulida (priapulid worms, from Gr. πριάπος,priāpos 'Priapus' + Lat.-ul-, diminutive), sometimes referred to aspenis worms, is aphylum of unsegmented marineworms. The name of the phylum relates to the Greek god of fertility, because their general shape and their extensible spiny introvert (eversible)proboscis may resemble the shape of a humanpenis. They live in the mud, except for a few tropical meiobenthic species which live in medium- to coarse-grained sands, and in comparatively shallow waters up to 90 metres (295 ft) deep and no warmer than 12–13°C.[3][4] Some species show a remarkable tolerance forhydrogen sulfide,anoxia and lowsalinity.[5][6]Halicryptus spinulosus appears to prefer brackish shallow waters.[7] They can be quite abundant in some areas. In an Alaskan bay as many as 85 adult individuals ofPriapulus caudatus per square meter has been recorded, while the density of its larvae can be as high as 58,000 per square meter (5,390 per square foot).[8]
Together withEchiura andSipuncula, they were once placed in the taxonGephyrea, but consistent morphological and molecular evidence supports their belonging toEcdysozoa, which also includesarthropods andnematodes. Fossil findings show that the mouth design of thestem-arthropodPambdelurion is identical with that of priapulids, indicating that their mouth is an original trait inherited from the last common ancestor of both priapulids and arthropods, even if modern arthropods no longer possess it.[9] Among Ecdysozoa, their nearest relatives areKinorhyncha andLoricifera, with which they constitute theScalidophora clade named after the spines covering the introvert (scalids).[10] They feed on slow-moving invertebrates, such aspolychaete worms.
Some analyses suggest that Priapulida may represent a basal lineage within Ecdysozoa, leading to their classification as "living fossils".[11] Priapulid-like fossils are known at least as far back as the MiddleCambrian. They were likely major predators of the Cambrian period. However, crown-group priapulids cannot be recognized until the Carboniferous.[1] 22 extant species of priapulid worms are known, half of them being ofmeiobenthic size.[12]
Priapulids are cylindrical worm-like animals, ranging from 0.2 to 0.3[13] to 39 centimetres[14] (0.08–0.12 to 15.35 in) long, with a median anterior mouth quite devoid of any armature or tentacles. They show both radial and bilateral symmetry. The gonads,protonephridia and ventral nerve cord are bilateral, while the introvert, pharynx and brain show radial symmetry, and appears to be a secondary trait.[15][16] Also the larvae show inernal and external characteristics of radial symmetry.[17] The adult body is divided into a main trunk or abdomen and a somewhat swollen proboscis region ornamented with longitudinal ridges. In addition it is ringed and often has circles of spines, which are continued into the slightly protrusible pharynx.[3] Family Priapulidae have species with a tail or a pair of caudal appendages. A slender tail or tail filament is also found in family Tubiluchidae. Appendages are absent in the remaining families.[18][19] The body has achitinous cuticle that is moulted as the animal grows.[20] Members of the family Chaetostephanidae also secretes a gelatinous tube, open in both ends, which they live in.[21]
Thealimentary canal is straight, consisting of an eversiblepharynx, an intestine, and a short rectum. The pharynx is muscular and lined by teeth.[20] Three of the five extant families have gone through a significant miniaturization and become detritivores (Tubiluchidae and Meiopriapulidae) and filter feeders (Chaetostephanidae). The two remaining families Priapulidae and Halicryptidae are larger carnivores that feed on other animals, although some species also consume detritus as larvae. The shape of the teeth reflect these different lifestyles, and seem to be adapted mainly towards grasping prey or raking detritus from the sediment into the mouth.[22][23] The anus is terminal, although inPriapulus one or two hollow ventral diverticula of the body-wall stretch out behind it.[3]
Thenervous system consists of a nerve ring around the pharynx and a prominent cord running the length of the body withganglia and longitudinal and transversal neurites consistent with an orthogonal organisation.[24] The nervous system retains abasiepidermal configuration with a connection with theectoderm, forming part of the body wall. There are no specializedsenseorgans, but there are sensory nerve endings in the body, especially on the proboscis.[20]
The priapulids aregonochoristic, having two separate sexes (i.e. male and female).[25] Their male and female organs are closely associated with the excretoryprotonephridia. They comprise a pair of branching tufts, each of which opens to the exterior on one side of the anus. The tips of these tufts enclose a flame-cell like those found inflatworms and other animals, and these probably function as excretory organs. As the animals mature, diverticula arise on the tubes of these organs, which develop either spermatozoa or ova. These sex cells pass out through the ducts.[3] The perigenital area of the genus Tubiluchus exhibit sexual dimorphism.[26]
For the speciesPriapulus caudatus, the 80 μm egg undergoes a total and radial cleavage following a symmetrical and subequal pattern.[27] Development is remarkably slow, with the first cleavage taking place 15 hours after fertilization, gastrulation after several days and hatching of the first 'lorica' larvae after 15 to 20 days.[28] The speciesMeiopriapulus fijiensis have direct development.[29] In current systematics, they are described as protostomes, despite having a deuterostomic development.[30] Because the group is so ancient, it is assumed the deuterostome condition which appears to be ancestral forbilaterians have been maintained.[31]
Stem-group priapulids are known from the Middle CambrianBurgess Shale, where their soft-part anatomy is preserved, often in conjunction with their gut contents – allowing a reconstruction of their diets.[32] In addition, isolatedmicrofossils (corresponding to the various teeth and spines that line the pharynx and introvert) are widespread in Cambrian deposits,[33] allowing the distribution of priapulids – and even individual species – to be tracked widely through Cambrian oceans.[34][22]Trace fossils that are morphologically almost identical to modern priapulid burrows (Treptichnus pedum) officially mark the start of theCambrian period, suggesting that priapulids, or at least close anatomical relatives, evolved around this time.[32] Crown-group priapulid body fossils are first known from the Carboniferous.[1]
^Oeschger, R.; Janssen, H. H. (September 1991). "Histological studies on Halicryptus spinulosus (Priapulida) with regard to environmental hydrogen sulfide resistance".Hydrobiologia.222:1–12.doi:10.1007/BF00017494.S2CID31342308.
^Dunn, C. W.; Hejnol, A.; Matus, D. Q.; Pang, K.; Browne, W. E.; Smith, S. A.; Seaver, E.; Rouse, G. W.; Obst, M. (10 April 2008). "Broad Phylogenomic Sampling Improves Resolution of the Animal Tree of Life".Nature.452 (7188):745–749.Bibcode:2008Natur.452..745D.doi:10.1038/nature06614.PMID18322464.S2CID4397099.
^Webster, Bonnie L.; Copley, Richard R.; Jenner, Ronald A.; Mackenzie-Dodds, Jacqueline A.; Bourlat, Sarah J.; Rota-Stabelli, Omar; Littlewood, D. T. J.; Telford, Maximilian J. (November 2006). "Mitogenomics and phylogenomics reveal priapulid worms as extant models of the ancestral Ecdysozoan".Evolution & Development.8 (6):502–510.doi:10.1111/j.1525-142x.2006.00123.x.
^Shirley, Thomas C.; Storch, Volker (1999). "Halicryptus higginsi n.sp. (Priapulida): A Giant New Species from Barrow, Alaska".Invertebrate Biology.118 (4):404–413.doi:10.2307/3227009.JSTOR3227009.
^Rothe, B. H.; Schmidt-Rhaesa, A. (Winter 2010). "Structure of the nervous system inTubiluchus troglodytes (Priapulida)".Invertebrate Biology.129:39–58.doi:10.1111/j.1744-7410.2010.00185.x.
^Pechenik, J. A. (2009).Biology of the Invertebrates (6th ed.). New York: McGraw-Hill. p. 454.ISBN978-0-07-302826-2.
^Higgins, Robert P.; Storch, Volker (January 1991). "Evidence for Direct Development in Meiopriapulus fijiensis (Priapulida)".Transactions of the American Microscopical Society.110 (1):37–46.doi:10.2307/3226738.JSTOR3226738.
One or more of the preceding sentences incorporates text from a publication now in thepublic domain: Shipley, Arthur Everett (1911). "Priapuloidea". InChisholm, Hugh (ed.).Encyclopædia Britannica. Vol. 22 (11th ed.). Cambridge University Press. p. 313.
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