Siboglinidae

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Siboglinidae
Temporal range: Pliensbachian-Holocene 189.6–Recent  Ma
Riftia tube worms Galapagos 2011.jpg
Riftia pachyptila
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Annelida
Clade: Pleistoannelida
Clade: Sedentaria
Infraclass: Canalipalpata
Order: Sabellida
Family: Siboglinidae
Caullery, 1914
Genera

See text

Siboglinidae is a family of polychaete annelid worms whose members made up the former phyla Pogonophora and Vestimentifera (the giant tube worms). [1] [2] The family is composed of around 100 species of vermiform creatures which live in thin tubes buried in sediment (Pogonophora) or in tubes attached to hard substratum (Vestimentifera) at ocean depths ranging from 100 to 10,000 m (300 to 32,800 ft). They can also be found in association with hydrothermal vents, methane seeps, sunken plant material, and whale carcasses.

Contents

The first specimen was dredged from the waters of Indonesia in 1900. These specimens were given to French zoologist Maurice Caullery, who studied them for nearly 50 years.

Anatomy

Most siboglinids are less than 1 millimetre (0.04 in) in diameter, but 10–75 centimetres (3.9–29.5 in) in length. They inhabit tubular structures composed of chitin which are fixed to rocks or substrates. The tubes are often clustered together in large colonies. [3]

Their bodies are divided into four regions. The anterior end is called the cephalic lobe, which ranges from one to over 200 thin branchial ciliated tentacles, each with tiny side branches known as pinnules. Behind this is a glandular forepart, which helps to secrete the tube. The main part of the body is the trunk, which is greatly elongated and bears various annuli, papillae, and ciliary tracts. Posterior to the trunk is the short metamerically segmented opisthosoma, bearing external paired chaetae, which help to anchor the animal to the base of its tube. [3]

The body cavity has a separate compartment in each of the first three regions of the body and extends into the tentacles. The opisthosoma has a coelomic chamber in each of its 5 to 23 segments, separated by septa. The worms have a complex closed circulatory system and a well-developed nervous system, but as adults, siboglinids completely lack a mouth, gut, and anus. [4]

Evolution

The family Siboglinidae has been difficult to place in an evolutionary context. [5] After examination of genetic differences between annelids, Siboglinidae were placed within the order Polychaeta by scientific consensus. [6] The fossil record along with molecular clocks suggest the family has Mesozoic (250 – 66 Mya) or Cenozoic (66 Mya – recent) origins. [5] However, some fossils of crystallized tubes are attributed to early Siboglinidae dating back to 500 Mya. [5] The oldest definitive specimens referred to the family came from Early Jurassic (Pliensbachian-Toarcian) Figueroa Sulfide deposits from San Rafael Mountains, found to be similar to modern Ridgeia . [7] This tubes, know as ‘Figueroa tubes’, along the ‘Troodos collared tubes’ (Cyprus, Turonian) were resolved among modern vestimentiferans. [8] Molecular work aligning five genes has identified four distinct clades within Siboglinidae. [9] [10] [11] The clades are Vestimentifera, Sclerolinum, Frenulata , and Osedax . [10] Vestimentiferans live in vent and seep habitats. [10] Separation of vestimentiferans into seep and deep-sea-dwelling clades is still debated due to some phylogenies based on sequencing data placing the genera along a continuum. [12] Sclerolinum is a monogeneric clade (which may be called Monilifera) living on organic-rich remains. [5] Frenulates live in organic-rich sediment habitats. [13] Osedax is a monogeneric clade specialized in living on whale bones, although recent evidence shows them living on fish bones as well. [14]

One probable relationship between the four clades is shown in the cladogram below. The position of Osedax is weakly supported. [5]

Siboglinidae

clade Frenulata

Osedax

Sclerolinum (clade Monilifera)

clade Vestimentifera

Vestimentiferans

Like other tube worms, vestimentiferans are benthic marine creatures. Riftia pachyptila , a vestimentiferan, is known only from the hydrothermal vent systems. [5]

Anatomy of vestimentiferans

Lamellibrachia satsuma removed from its tube: op = opisthosome, ves = vestimentum, ten = tentacular region, tr = trunk Lamellibrachia satsuma.png
Lamellibrachia satsuma removed from its tube: op = opisthosome, ves = vestimentum, ten = tentacular region, tr = trunk

Vestimentiferan bodies are divided into four regions: the obturaculum, vestimentum, trunk, and opisthosome. The main trunk of the body bears wing-like extensions. Unlike other siboglinids that never have a digestive tract, they have one that they completely lose during metamorphosis.

The obturaculum is the first anterior body part. [15] It is possible that the obturaculum is actually an outgrowth of the vestimentum rather than a separate body segment which would distinguish it from other siboglinids.

The vestimentum, from which the group's name is derived, is a wing-like body part with glands that secrete the tube. In a ventroanterior position in the vestimentum is the brain which is postulated to be simpler than relatives that maintain a gut in the adult form. [15] The opisthosome is the anchoring rear body part.

Vestimentiferan ecology

Their primary nutrition is derived from the sulfide-rich fluids emanating from the hydrothermal vents where they live. The sulfides are metabolized by symbiotic hydrogen sulfide- or methane-oxidizing bacteria living in an internal organ, the trophosome. One gram of trophosome tissue can contain one billion bacteria. The origin of this symbiotic relationship is not currently known. The bacteria appear to colonize the host animal larvae after they have settled on a surface, entering them through their skin. [16] This method of entry, known as horizontal transmission, means that each organism may have different species of bacteria assisting in this symbiosis. However, these bacteria all play similar roles in sustaining the vestimentiferans. Endosymbionts have a wide variety of metabolic genes, which may allow them to switch between autotrophic and heterotrophic methods of nutrient acquisition. [17] When the host dies, the bacteria are released and return to the free-living population in the seawater. [18]

Discovery of the hydrothermal vents in the eastern Pacific Ocean was quickly followed by the discovery and description of new vestimentiferan tubeworm species. These tubeworms are one of the most dominant organisms associated with the hydrothermal vents in the Pacific Ocean. Tubeworms anchor themselves to the substratum of the hydrocarbon seep by roots located at the basal portion of their bodies. [19] Intact tubeworm roots have proven very difficult to obtain for study because they are extremely delicate, and often break off when a tubeworm is removed from hypothermal vent regions. How long the roots of the tube worms can grow is unknown, but roots have been recovered longer than 30 m.[ citation needed ]

A single aggregation of tubeworms can contain thousands of individuals, and the roots produced by each tubeworm can become tangled with the roots of neighbouring tubeworms. [20] These mats of roots are known as "ropes", and travel down the tubes of dead tubeworms, and run through holes in rocks. The diameter and wall thickness of the tubeworm roots do not appear to change with distance from the trunk portion of the tubeworm's body.

Like the trunk portion of the body, the roots of the vestimentiferan tubeworms are composed of chitin crystallites, which support and protect the tubeworm from predation and environmental stresses. Tubeworms build the external chitin structure themselves by secreting chitin from specialized glands located in their body walls.

Genera

Related Research Articles

<span class="mw-page-title-main">Marine worm</span>

Any worm that lives in a marine environment is considered a water worm. Marine worms are found in several different phyla, including the Platyhelminthes, Nematoda, Annelida, Chaetognatha, Hemichordata, and Phoronida. For a list of marine animals that have been called "sea worms", see sea worm.

<span class="mw-page-title-main">Polychaete</span> Class of annelid worms

Polychaeta is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes. Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made of chitin. More than 10,000 species are described in this class. Common representatives include the lugworm and the sandworm or clam worm Alitta.

<span class="mw-page-title-main">Cold seep</span> Ocean floor area where hydrogen sulfide, methane and other hydrocarbon-rich fluid seepage occurs

A cold seep is an area of the ocean floor where seepage of fluids rich in hydrogen sulfide, methane, and other hydrocarbons occurs, often in the form of a brine pool. Cold does not mean that the temperature of the seepage is lower than that of the surrounding sea water; on the contrary, its temperature is often slightly higher. The "cold" is relative to the very warm conditions of a hydrothermal vent. Cold seeps constitute a biome supporting several endemic species.

<i>Riftia</i> Giant tube worm (species of annelid)

Riftia pachyptila, commonly known as the giant tube worm and less commonly known as the giant beardworm, is a marine invertebrate in the phylum Annelida related to tube worms commonly found in the intertidal and pelagic zones. R. pachyptila lives on the floor of the Pacific Ocean near hydrothermal vents. The vents provide a natural ambient temperature in their environment ranging from 2 to 30 °C, and this organism can tolerate extremely high hydrogen sulfide levels. These worms can reach a length of 3 m, and their tubular bodies have a diameter of 4 cm (1.6 in).

<i>Osedax</i> Genus of annelid worms

Osedax is a genus of deep-sea siboglinid polychaetes, commonly called boneworms, zombie worms, or bone-eating worms. Osedax is Latin for "bone-eater". The name alludes to how the worms bore into the bones of whale carcasses to reach enclosed lipids, on which they rely for sustenance. They utilize specialized root tissues for bone-boring. It is possible that multiple species of Osedax reside in the same bone. Osedax worms are also known to feed on the collagen itself by making holes in the whale's skeletal structure. These holes can also serve as a form of protection from nearby predators.

<i>Lamellibrachia</i> Genus of annelids

Lamellibrachia is a genus of tube worms related to the giant tube worm, Riftia pachyptila. They live at deep-sea cold seeps where hydrocarbons leak out of the seafloor, and are entirely reliant on internal, sulfide-oxidizing bacterial symbionts for their nutrition. The symbionts, gammaproteobacteria, require sulfide and inorganic carbon. The tube worms extract dissolved oxygen and hydrogen sulfide from the sea water with the crown of plumes. Species living near seeps can also obtain sulfide through their "roots", posterior extensions of their body and tube. Several sorts of hemoglobin are present in the blood and coelomic fluid to bind to the different components and transport them to the symbionts.

<span class="mw-page-title-main">Metatrochophore</span>

A metatrochophore (;) is a type of larva developed from the trochophore larva of a polychaete annelid.

<i>Serpula</i> Genus of annelid worms

Serpula is a genus of sessile, marine annelid tube worms that belongs to the family Serpulidae. Serpulid worms are very similar to tube worms of the closely related sabellid family, except that the former possess a cartilaginous operculum that occludes the entrance to their protective tube after the animal has withdrawn into it. The most distinctive feature of worms of the genus Serpula is their colorful fan-shaped "crown". The crown, used by these animals for respiration and alimentation, is the structure that is most commonly seen by scuba divers and other casual observers.

<i>Lamellibrachia luymesi</i> Species of tube worms in the family Siboglinidae

Lamellibrachia luymesi is a species of tube worms in the family Siboglinidae. It lives at deep-sea cold seeps where hydrocarbons are leaking out of the seafloor. It is entirely reliant on internal, sulfide-oxidizing bacterial symbionts for its nutrition. These are located in a centrally located "trophosome".

<span class="mw-page-title-main">Trophosome</span> Organ containing endosymbionts

A trophosome is a highly vascularised organ found in some animals that houses symbiotic bacteria that provide food for their host. Trophosomes are contained by the coelom of tube worms and in the body of symbiotic flatworms of the genus Paracatenula.

<span class="mw-page-title-main">Errantia</span> Subclass of annelid worms

Errantia is a diverse group of marine polychaete worms in the phylum Annelida. Traditionally a subclass of the paraphyletic class Polychaeta, it is currently regarded as a monophyletic group within the larger Pleistoannelida, composed of Errantia and Sedentaria. These worms are found worldwide in marine environments and brackish water.

Osedax japonicus is a species of bathypelagic polychaete tube worm that lives at great depths on the seabed and is able to sustain itself on the bones of a dead whale. It was first described in 2006 from a sunken sperm whale carcase near Kyushu, Japan.

<span class="mw-page-title-main">Annelid</span> Phylum of segmented worms

The annelids, also known as the segmented worms, comprise a large phylum called Annelida. It contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecologies – some in marine environments as distinct as tidal zones and hydrothermal vents, others in fresh water, and yet others in moist terrestrial environments.

<span class="mw-page-title-main">Marine microbial symbiosis</span>

Microbial symbiosis in marine animals was not discovered until 1981. In the time following, symbiotic relationships between marine invertebrates and chemoautotrophic bacteria have been found in a variety of ecosystems, ranging from shallow coastal waters to deep-sea hydrothermal vents. Symbiosis is a way for marine organisms to find creative ways to survive in a very dynamic environment. They are different in relation to how dependent the organisms are on each other or how they are associated. It is also considered a selective force behind evolution in some scientific aspects. The symbiotic relationships of organisms has the ability to change behavior, morphology and metabolic pathways. With increased recognition and research, new terminology also arises, such as holobiont, which the relationship between a host and its symbionts as one grouping. Many scientists will look at the hologenome, which is the combined genetic information of the host and its symbionts. These terms are more commonly used to describe microbial symbionts.

<i>Tevnia</i> Genus of annelid worms

Tevnia is a genus of giant tube worm in the family Siboglinidae, with only one species, Tevnia jerichonana, living in a unique deep-sea environment. These deep sea marine species survive in environments like hydrothermal vents. These vents give off gas and toxic chemicals with the addition of having superheated temperatures. The giant tube worm prefers environments such as these despite the harsh temperature and toxic sea water.

Hydrogen sulfide chemosynthesis is a form of chemosynthesis which uses hydrogen sulfide. It is common in hydrothermal vent microbial communities Due to the lack of light in these environments this is predominant over photosynthesis

<i>Lamellibrachia satsuma</i> Species of tube worms in the family Siboglinidae

Lamellibrachia satsuma is a vestimentiferan tube worm that was discovered near a hydrothermal vent in Kagoshima Bay, Kagoshima at the depth of only 82 m (269 ft) the shallowest depth record for a vestimentiferan. Its symbiotic sulfur oxidizer bacteria have been characterised as ε-Proteobacteria and γ-Proteobacteria. Subspecies have been later found associated with cold seeps at Hatsushima in Sagami Bay and at the Daini Tenryu Knoll in the Nankai Trough with specimens obtained at up to 1,170 m (3,840 ft) depth.

<span class="mw-page-title-main">Pleistoannelida</span> Clade of annelid worms

Pleistoannelida is a group of annelid worms that comprises the vast majority of the diversity in phylum Annelida. Discovered through phylogenetic analyses, it is the largest clade of annelids, comprised by the last common ancestor of the highly diverse sister groups Errantia and Sedentaria and all the descendants of that ancestor. Most groups in the Clade find their ancestors within the Cambrian explosion when Annelid diversity expanded dramatically. The Pleistoannelida clade covers a variety of traits. However, the evolution of simple to complex eyes, developed papillae for burrowing, and for some specialized radioles for feeding can be seen universally across every species. New findings have discovered the range of Annelid diversity have led to uncertainty if groups with developed ancestral traits should remain within the clade. Furthermore, there's been a lack of recently discovered Annelid traits being used in the categorization of groups within the clade, leading to many hypothesis on how to do so and which should remain within the clade. Currently three smaller clades that were originally a part of the groups Errantia and Sedentaria have been proven to fall outside while still being connected to the basal groups.

Oligobrachia is a genus in the family Siboglinidae, commonly known as beard worms. These beard worms are typically found at spreading centers, hydrothermal vents, and undersea volcanoes. The siboglinidae are annelids which can be found buried in sediments. Beard worms do not necessarily exist at one specific part of the world's oceans, however, they are spread out all over the ocean floors as long as the surrounding environment is similar; these are known as metapopulations. Most commonly, these organisms are found at the bottom of the ocean floor, whether it be at a depth of roughly 25 meters or hundreds of meters. Oligobrachia can typically be found near hydrothermal vents and methane seeps. An important characteristic of this genus is that it lacks a mouth and gut. Therefore, it relies on symbiotic bacteria to provide the host organism with energy to survive. The majority of oligobrachia that have been observed have been found in the Arctic and other high-latitude areas of the world's oceans.

Frenulata, "beard worms", is a clade of Siboglinidae, "tube worms". They are one of four lineages with numerous species. They may be the most basal clade in the family. Despite being the first tube worms to be encountered and described, they remain the least studied group. This is because of their slender shape, they often get destroyed as a result of being caught as bycatch or poor preservation. They are found primarily in deep, muddy sediments, cold seeps, and anoxic firth sediments.

References

  1. ↑ Kojima, S.; Hashimoto, T.; Hasegawa, M.; Murata, S.; Ohta, S.; Seki, H.; Okada, N. (July 1993). "Close phylogenetic relationship between vestimentifera (tube worms) and annelida revealed by the amino acid sequence of elongation factor-lα". Journal of Molecular Evolution. 37 (1): 66–70. Bibcode:1993JMolE..37...66K. doi:10.1007/BF00170463. PMID   8360920. S2CID   23391565.
  2. 1 2 3 4 Rouse, G. W. (2001). "A cladistic analysis of Siboglinidae Caullery, 1914 (Polychaeta, Annelida): formerly the phyla Pogonophora and Vestimentifera". Zoological Journal of the Linnean Society. 132 (1): 55–80. doi: 10.1006/zjls.2000.0263 .
  3. 1 2 Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 862–863. ISBN   0-03-056747-5.
  4. ↑ Ross Piper (30 August 2007). Extraordinary animals: an encyclopedia of curious and unusual animals . Greenwood Publishing Group. ISBN   978-0-313-33922-6 . Retrieved 3 December 2011.
  5. 1 2 3 4 5 6 Hilário, Ana; Capa, María; Dahlgren, Thomas G.; Halanych, Kenneth M.; Little, Crispin T. S.; Thornhill, Daniel J.; Verna, Caroline; Glover, Adrian G. (2011). Laudet, Vincent (ed.). "New Perspectives on the Ecology and Evolution of Siboglinid Tubeworms". PLOS ONE. 6 (2): e16309. Bibcode:2011PLoSO...616309H. doi: 10.1371/journal.pone.0016309 . PMC   3038861 . PMID   21339826.
  6. ↑ Struck, T. H.; Schult, N.; Kusen, T.; Hickman, E.; Bleidorn, C.; McHugh, D.; Halanych, K. M. (2007). "Annelid phylogeny and the status of Sipuncula and Echiura". BMC Evolutionary Biology. 7: 57. doi: 10.1186/1471-2148-7-57 . PMC   1855331 . PMID   17411434.
  7. ↑ LITTLE, CRISPIN T. S.; DANELIAN, TANIEL; HERRINGTON, RICHARD J.; HAYMON, RACHEL M. (2004). "Early jurassic hydrothermal vent community from the Franciscan Complex, California". Journal of Paleontology. 78 (3): 542–559. doi:10.1666/0022-3360(2004)078<0542:ejhvcf>2.0.co;2. ISSN   0022-3360.
  8. ↑ Georgieva, Magdalena N.; Little, Crispin T. S.; Watson, Jonathan S.; Sephton, Mark A.; Ball, Alexander D.; Glover, Adrian G. (2017-12-28). "Identification of fossil worm tubes from Phanerozoic hydrothermal vents and cold seeps". Journal of Systematic Palaeontology. 17 (4): 287–329. doi:10.1080/14772019.2017.1412362. ISSN   1477-2019.
  9. ↑ Black, M. B.; Halanych, K. M.; Maas, P. A. Y.; Hoeh, W. R.; Hashimoto, J.; Desbruyeres, D.; Lutz, R. A.; et al. (1997). "Molecular systematics of vestimentiferan tubeworms from hydrothermal vents and cold-water seeps". Marine Biology. 130 (2): 141–149. doi:10.1007/s002270050233. S2CID   13217485.
  10. 1 2 3 Glover, A. G.; Kallstrom, B.; Smith, C. R.; Dahlgren, T. G. (2005). "World-wide whale worms? A new species of Osedax from the shallow north Atlantic". Proceedings of the Royal Society B: Biological Sciences. 272 (1581): 2587–2592. doi:10.1098/rspb.2005.3275. PMC   1559975 . PMID   16321780.
  11. ↑ Vrijenhoek, R. C.; Johnson, S. B.; Rouse, G. W. (2009). "A remarkable diversity of bone-eating worms (Osedax; Siboglinidae; Annelida)". BMC Biology. 7: 74. doi: 10.1186/1741-7007-7-74 . PMC   2780999 . PMID   19903327.
  12. ↑ Bright, Monika; Lallie, François (2010-05-12), Gibson, R; Atkinson, R; Gordon, J (eds.), "The Biology of Vestimentiferan Tubeworms", Oceanography and Marine Biology, vol. 20103650, CRC Press, pp. 213–265, doi:10.1201/ebk1439821169-c4, ISBN   9781439821169
  13. ↑ Rodrigues, C. F.; Hilário, A.; Cunha, M. R.; Weightman, A. J.; Webster, G. (2011). "Microbial diversity in Frenulata (Siboglinidae, Polychaeta) species from mud volcanoes in the Gulf of Cadiz (NE Atlantic)". Antonie van Leeuwenhoek. 100 (1): 83–98. doi:10.1007/s10482-011-9567-0. PMID   21359663. S2CID   10224623.
  14. ↑ Rouse, G. W.; Goffredi, S. K.; Johnson, S. B.; Vrijenhoek, R. C. (2011). "Not whale-fall specialists, Osedax worms also consume fishbones". Biology Letters. 7 (5): 736–739. doi:10.1098/rsbl.2011.0202. PMC   3169056 . PMID   21490008.
  15. 1 2 Miyamoto, Norio; Shinozaki, Ayuta; Fujiwara, Yoshihiro (2013-01-23). "Neuroanatomy of the Vestimentiferan Tubeworm Lamellibrachia satsuma Provides Insights into the Evolution of the Polychaete Nervous System". PLOS ONE. 8 (1): e55151. doi: 10.1371/journal.pone.0055151 . PMC   3553155 . PMID   23372830.
  16. ↑ Nussbaumer, Andrea D.; Fisher, Charles R.; Bright, Monika (2006-05-18). "Horizontal endosymbiont transmission in hydrothermal vent tubeworms". Nature. 441 (7091): 345–348. Bibcode:2006Natur.441..345N. doi:10.1038/nature04793. ISSN   1476-4687. PMID   16710420. S2CID   18356960.
  17. ↑ Reveillaud, Julie; Anderson, Rika; Reves-Sohn, Sintra; Cavanaugh, Colleen; Huber, Julie A. (2018-01-27). "Metagenomic investigation of vestimentiferan tubeworm endosymbionts from Mid-Cayman Rise reveals new insights into metabolism and diversity". Microbiome. 6 (1): 19. doi: 10.1186/s40168-018-0411-x . ISSN   2049-2618. PMC   5787263 . PMID   29374496.
  18. ↑ Klose, Julia; Polz, Martin F.; Wagner, Michael; Schimak, Mario P.; Gollner, Sabine; Bright, Monika (2015-09-08). "Endosymbionts escape dead hydrothermal vent tubeworms to enrich the free-living population". Proceedings of the National Academy of Sciences of the United States of America. 112 (36): 11300–11305. Bibcode:2015PNAS..11211300K. doi: 10.1073/pnas.1501160112 . ISSN   1091-6490. PMC   4568656 . PMID   26283348.
  19. ↑ Halanych, K.; Maas, P.; Hoeh, W.; Hashimoto, J.; Desbruyeres, D.; Lutz, R.; Vrijenhoek, R. (1997). "Molecular systematics of vestimentiferan tubeworms from hydrothermal vents and cold-water seeps". Marine Biology. 130 (2): 141–149. doi:10.1007/s002270050233. S2CID   13217485.
  20. ↑ Julian, D.; Gaill, F.; Wood, E.; Arp, A.; Fisher, C. (1999). "Roots as a site of hydrogen sulphide uptake in the hydrocarbon seep vestimentiferan Lamellibrachia sp". The Journal of Experimental Biology. 202 (Pt 17): 2245–57. doi:10.1242/jeb.202.17.2245. PMID   10441078.
  21. ↑ Hilário, A.; Cunha, M. R. (2008). "On some frenulate species (Annelida: Polychaeta: Siboglinidae) from mud volcanoes in the Gulf of Cadiz (NE Atlantic)". Scientia Marina. 72 (2): 361–371. doi: 10.3989/scimar.2008.72n2361 .
  22. ↑ Southward, E. C.; Schulze, A.; Tunnicliffe, V. (2002). "Vestimentiferans (Pogonophora) in the Pacific and Indian Oceans: a new genus from Lihir Island (Papua New Guinea) and the Java Trench, with the first report of Arcovestia ivanovi from the North Fiji Basin". Journal of Natural History. 36 (10): 1179–1197. doi:10.1080/00222930110040402. S2CID   86076917.
  23. ↑ Smirnov, R. V. (2000). "A redescription of Spirobrachia leospira Gureeva (Pogonophora), with the erection of a new genus and a revision of the Spirobrachiidae". Ophelia. 53 (2): 151–158. doi:10.1080/00785236.2000.10409445. S2CID   85319259.
  24. ↑ Halanych, Kenneth M. (2005). "Molecular phylogeny of siboglinid annelids (a.k.a. pogonophorans): a review". Hydrobiologia. 535–536 (1): 297–307. doi:10.1007/s10750-004-1437-6. ISSN   0018-8158. S2CID   16022909.
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