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{{Short description|Family of annelid worms}}{{Automatic taxobox
{{Taxobox
| fossil_range = {{Fossil range|189.6|Recent|earliest=500|latest=0}}
⚫
| name = Siboglinidae
| image = Riftia tube worms Galapagos 2011.jpg
| image = Riftia tube worms Galapagos 2011.jpg
| image_width = 250px
| image_upright = 1.2
| image_caption = ''[[Riftia pachyptila]]''
| image_caption = ''[[Riftia pachyptila]]''
| display_parents = 2
| regnum = [[Animal]]ia
⚫
| taxon = Siboglinidae
| subregnum = [[Eumetazoa]]
⚫
| authority = [[Maurice Caullery|Caullery]], 1914
| phylum = [[Annelid]]a
⚫
| subdivision_ranks = Genera
| classis = [[Polychaete|Polychaeta]]
⚫
| subdivision = [[Siboglinidae#Genera|See text]]
| subclassis = [[Palpata]]
| ordo = [[Canalipalpata]]
⚫
| familia = '''Siboglinidae'''
⚫
| familia_authority = [[Maurice Caullery|Caullery]], 1914
⚫
| subdivision = See text.
⚫
| subdivision_ranks = [[Genus|Genera]]
}}
}}


'''Siboglinidae''', also known as the '''beard worms''', is a [[family (biology)|family]] of [[polychaete]] [[Annelida|annelid worms]] whose members made up the former [[phylum|phyla]] '''Pogonophora''' (the [[giant tube worms]]) and '''Vestimentifera'''.<ref>{{cite journal | title=Close phylogenetic relationship between vestimentifera (tube worms) and annelida revealed by the amino acid sequence of elongation factor-lα | doi=10.1007/BF00170463 | journal=Journal of Molecular Evolution | volume=37 | issue=1 |date=July 1993 | author=Shigeaki Kojima, Tetsuo Hashimoto, Masami Hasegawa, Shigenori Murata, Suguru Ohta, Humitake Seki and Norihiro Okada | pages=66–70 | pmid=8360920}}</ref><ref name=Rouse>{{cite journal | author=Rouse G.W. |title=A cladistic analysis of Siboglinidae Caullery, 1914 (Polychaeta, Annelida): formerly the phyla Pogonophora and Vestimentifera | journal=Zoological Journal of the Linnean Society | volume=132 | issue=1| year=2001 | pages=55–80 | doi = 10.1006/zjls.2000.0263}}</ref> They are composed of about 100 species of vermiform creatures and live in thin tubes buried in sediments at ocean depths from 100 to 10,000&nbsp;m. They can also be found in association with [[hydrothermal vent]]s, [[cold seep|methane seep]]s, with sunken plant material or [[Whale fall|whale carcasses]].
'''Siboglinidae''' is a [[family (biology)|family]] of [[polychaete]] [[Annelida|annelid worms]] whose members made up the former [[phylum|phyla]] '''Pogonophora''' and '''Vestimentifera''' (the [[giant tube worm]]s).<ref>{{cite journal | title=Close phylogenetic relationship between vestimentifera (tube worms) and annelida revealed by the amino acid sequence of elongation factor-lα | doi=10.1007/BF00170463 | journal=Journal of Molecular Evolution | volume=37 | issue=1 |date=July 1993 | last1=Kojima |first1=S. |last2=Hashimoto |first2=T. |last3=Hasegawa |first3=M. |last4=Murata |first4=S. |last5=Ohta |first5=S. |last6=Seki |first6=H. |last7=Okada |first7=N. | pages=66–70 | pmid=8360920| bibcode=1993JMolE..37...66K | s2cid=23391565 }}</ref><ref name=Rouse>{{cite journal | last=Rouse | first=G. W. |title=A cladistic analysis of Siboglinidae Caullery, 1914 (Polychaeta, Annelida): formerly the phyla Pogonophora and Vestimentifera | journal=Zoological Journal of the Linnean Society | volume=132 | issue=1| year=2001 | pages=55–80 | doi = 10.1006/zjls.2000.0263| doi-access=free }}</ref> 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 {{convert|100|to|10000|m|ft|abbr=on|-2}}. They can also be found in association with [[hydrothermal vent]]s, [[methane seep]]s, sunken plant material, and [[Whale fall|whale carcasses]].


The first specimen was dredged from the waters of what is now [[Indonesia]] in 1900. These specimens were given to French zoologist [[Maurice Caullery]], who studied them for nearly 50&nbsp;years.
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==
==Anatomy==
Most siboglinids are less than {{convert|1|mm|in|sigfig=1}} in diameter but {{convert|10|-|75|cm|in}} in length. They inhabit tubular structures composed of [[chitin]] and fixed to the bottom. The tubes are often clustered together in large colonies.<ref name=IZ>{{cite book |author= Barnes, Robert D. |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia, PA|pages= 862–863|isbn= 0-03-056747-5}}</ref>
Most siboglinids are less than {{convert|1|mm|in|sigfig=1}} in diameter, but {{convert|10|-|75|cm|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.<ref name=IZ>{{cite book |author= Barnes, Robert D. |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia, PA|pages= 862–863|isbn= 0-03-056747-5}}</ref>


The body is divided into four regions. The anterior end is called the cephalic lobe, which bears from 1 to over 200 thin branchial [[cilia]]ted [[tentacle]]s, each bearing 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 [[Metamerism (biology)|metamerically]] segmented [[opisthosoma]], bearing external paired [[chaeta]]e, which apparently help to anchor the animal to the base of its tube.<ref name=IZ/>
Their bodies are divided into four regions. The anterior end is called the cephalic lobe, which ranges from one to over 200 thin branchial [[cilia]]ted [[tentacle]]s, 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 [[Metamerism (biology)|metamerically]] segmented [[opisthosoma]], bearing external paired [[chaeta]]e, which help to anchor the animal to the base of its tube.<ref name=IZ/>


The body cavity has a separate compartment in each of the first three regions of the body, and extends into the tentacles. The opisthoma has a [[coelom]]ic chamber in each of its 5 to 23 segments, separated by [[septum|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.<ref name=Extra>{{cite book|author=[[Ross Piper]]|title=Extraordinary animals: an encyclopedia of curious and unusual animals|url=http://books.google.com/books?id=eqegRf2UstIC|accessdate=3 December 2011|date=30 August 2007|publisher=Greenwood Publishing Group|isbn=978-0-313-33922-6}}</ref>
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 [[coelom]]ic chamber in each of its 5 to 23 segments, separated by [[septum|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.<ref name=Extra>{{cite book|author=Ross Piper|title=Extraordinary animals: an encyclopedia of curious and unusual animals|url=https://archive.org/details/extraordinaryani0000pipe|url-access=registration|access-date=3 December 2011|date=30 August 2007|publisher=Greenwood Publishing Group|isbn=978-0-313-33922-6|author-link=Ross Piper}}</ref>

Siboglinids are [[dioecious]], with one [[gonad]] on each side of the trunk, within the body cavity. The fertilised eggs develop within the tubes, and hatch to produce small ciliated worm-like larvae.<ref name=IZ/>


==Evolution==
==Evolution==
The family Siboglinidae has been difficult to place in an evolutionary context.<ref name="ReferenceA">{{cite journal|doi=10.1371/journal.pone.0016309|title=New Perspectives on the Ecology and Evolution of Siboglinid Tubeworms|year=2011|editor1-last=Laudet|editor1-first=Vincent|last1=Hilário|first1=Ana|last2=Capa|first2=María|last3=Dahlgren|first3=Thomas G.|last4=Halanych|first4=Kenneth M.|last5=Little|first5=Crispin T. S.|last6=Thornhill|first6=Daniel J.|last7=Verna|first7=Caroline|last8=Glover|first8=Adrian G.|journal=PLoS ONE|volume=6|issue=2|pages=e16309|pmid=21339826|pmc=3038861}}</ref> By examining genetic differences between annelids there is now a scientific consensus that Siboglinidae is a family within the order Polychaeta.<ref>{{cite journal|doi=10.1186/1471-2148-7-57|title=Annelid phylogeny and the status of Sipuncula and Echiura|year=2007|author=Struck, T. H., Schult, N., Kusen, T., Hickman, E., Bleidorn, C., McHugh, D., and Halanych, K. M.|journal=BMC Evol Biol|volume=7|pages=57|pmid=17411434|pmc=1855331}}</ref> The fossil record along with molecular clocks suggests the family has Mesozoic (250 mya - 66 mya) or Cenozoic (66 mya - recent) origins.<ref name="ReferenceA"/> Examination of molecular work aligning five genes has made clear that there are four distinct clades within Siboglinidae.<ref>{{cite journal|doi=10.1007/s002270050233|title=Molecular systematics of vestimentiferan tubeworms from hydrothermal vents and cold-water seeps|year=1997|author=Black, M. B., K. M. Halanych, P. A. Y. Maas, W. R. Hoeh, J. Hashimoto, D. Desbruyeres, R. A. Lutz|journal=Marine Biology|volume=130|issue=2|pages=141–149|display-authors=etal}}</ref><ref name=Glover>{{cite journal|doi=10.1098/rspb.2005.3275|title=World-wide whale worms? A new species of Osedax from the shallow north Atlantic|year=2005|author=Glover, A. G., B. Kallstrom, C. R. Smith, and T. G. Dahlgren|journal=Proceedings of the Royal Society B-Biological Sciences|volume=272|issue=1581|pages=2587–2592|pmid=16321780|pmc=1559975}}</ref><ref>{{cite journal|doi=10.1186/1741-7007-7-74|title=A remarkable diversity of bone-eating worms (Osedax; Siboglinidae; Annelida)|year=2009|author=Vrijenhoek, R. C., S. B. Johnson, and G. W. Rouse|journal=BMC Biol|volume=7|pages=74|pmid=19903327|pmc=2780999}}</ref> The clades are vestimentiferans, ''Sclerolinum'', frenulates, and ''Osedax''.<ref name=Glover/> These clades represent the four tracks evolution has followed within Siboglinidae.<ref name=Glover/> Vestimentiferans live in vent and seep habitats.<ref name=Glover/> ''Sclerolinum'' is a monogeneric clade living on organic-rich remains.<ref name="ReferenceA"/> Frenulates live in organic-rich sediment habitats.<ref>{{cite journal|doi=10.1007/s10482-011-9567-0|title=Microbial diversity in Frenulata (Siboglinidae, Polychaeta) species from mud volcanoes in the Gulf of Cadiz (NE Atlantic)|year=2011|author=Rodrigues, C. F., Hilario, A., Cunha, M. R., Weightman, A. J., and Webster, G.|journal=Antonie Van Leeuwenhoek|volume=100|pages=83–98}}</ref> ''Osedax'' is a monogeneric clade specialized in living on whale bones, although recently there has been evidence of them living on fish bones as well.<ref>{{cite journal|doi=10.1098/rsbl.2011.0202|title=Not whale-fall specialists, Osedax worms also consume fishbones|year=2011|author=Rouse, G. W., Goffredi, S. K., Johnson, S. B., and Vrijenhoek, R. C.|journal=Biology Letters|volume=7|issue=5|pages=736–739}}</ref>
The family Siboglinidae has been difficult to place in an evolutionary context.<ref name="ReferenceA">{{cite journal|doi=10.1371/journal.pone.0016309|title=New Perspectives on the Ecology and Evolution of Siboglinid Tubeworms|year=2011|editor1-last=Laudet|editor1-first=Vincent|last1=Hilário|first1=Ana|last2=Capa|first2=María|last3=Dahlgren|first3=Thomas G.|last4=Halanych|first4=Kenneth M.|last5=Little|first5=Crispin T. S.|last6=Thornhill|first6=Daniel J.|last7=Verna|first7=Caroline|last8=Glover|first8=Adrian G.|journal=PLOS ONE|volume=6|issue=2|pages=e16309|pmid=21339826|pmc=3038861|bibcode=2011PLoSO...616309H|doi-access=free}}</ref> After examination of genetic differences between annelids, Siboglinidae were placed within the order [[Polychaeta]] by scientific consensus.<ref>{{cite journal|doi=10.1186/1471-2148-7-57|title=Annelid phylogeny and the status of Sipuncula and Echiura|year=2007 |last1=Struck |first1=T. H. |last2=Schult |first2=N. |last3=Kusen |first3=T. |last4=Hickman |first4=E. |last5=Bleidorn |first5=C. |last6=McHugh |first6=D. |last7=Halanych |first7=K. M. |journal=BMC Evolutionary Biology |volume=7|pages=57|pmid=17411434|pmc=1855331 |doi-access=free }}</ref> The fossil record along with [[molecular clock]]s suggest the family has [[Mesozoic]] (250 – 66 Mya) or [[Cenozoic]] (66 Mya – recent) origins.<ref name="ReferenceA" /> However, some fossils of crystallized tubes are attributed to early Siboglinidae dating back to 500 Mya.<ref name="ReferenceA" /> Molecular work aligning five genes has identified four distinct [[clade]]s within Siboglinidae.<ref>{{cite journal|doi=10.1007/s002270050233|title=Molecular systematics of vestimentiferan tubeworms from hydrothermal vents and cold-water seeps|year=1997 |last1=Black |first1=M. B. |first2=K. M. |last2=Halanych |first3=P. A. Y. |last3=Maas |first4=W. R. |last4=Hoeh |first5=J. |last5=Hashimoto |first6=D. |last6=Desbruyeres |first7=R. A. |last7=Lutz |journal=Marine Biology|volume=130|issue=2|pages=141–149|s2cid=13217485|display-authors=etal}}</ref><ref name="Glover">{{cite journal|doi=10.1098/rspb.2005.3275|title=World-wide whale worms? A new species of Osedax from the shallow north Atlantic|year=2005 |last1=Glover |first1=A. G. |first2=B. |last2=Kallstrom |first3=C. R. |last3=Smith |first4=T. G. |last4=Dahlgren |journal=Proceedings of the Royal Society B: Biological Sciences|volume=272|issue=1581|pages=2587–2592|pmid=16321780|pmc=1559975}}</ref><ref>{{cite journal|doi=10.1186/1741-7007-7-74|title=A remarkable diversity of bone-eating worms (Osedax; Siboglinidae; Annelida)|year=2009 |last1=Vrijenhoek |first1=R. C. |first2=S. B. |last2=Johnson |first3=G. W. |last3=Rouse|journal=BMC Biology |volume=7|pages=74|pmid=19903327|pmc=2780999 |doi-access=free }}</ref> The clades are ''Vestimentifera'', ''Sclerolinum'', ''[[Frenulata]]'', and ''[[Osedax]]''.<ref name=Glover/> Vestimentiferans live in vent and seep habitats.<ref name=Glover/> 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.<ref>{{Citation|last1=Bright|first1=Monika|title=The Biology of Vestimentiferan Tubeworms|date=2010-05-12|work=Oceanography and Marine Biology|volume=20103650|pages=213–265|editor-last=Gibson|editor-first=R|publisher=CRC Press|language=en|doi=10.1201/ebk1439821169-c4|isbn=9781439821169|last2=Lallie|first2=François|editor2-last=Atkinson|editor2-first=R|editor3-last=Gordon|editor3-first=J}}</ref> ''Sclerolinum'' is a monogeneric clade (which may be called Monilifera) living on organic-rich remains.<ref name="ReferenceA"/> Frenulates live in organic-rich sediment habitats.<ref>{{cite journal|doi=10.1007/s10482-011-9567-0|pmid=21359663|title=Microbial diversity in Frenulata (Siboglinidae, Polychaeta) species from mud volcanoes in the Gulf of Cadiz (NE Atlantic)|year=2011 |last1=Rodrigues |first1=C. F. |last2=Hilário |first2=A. |last3=Cunha |first3=M. R. |last4=Weightman |first4=A. J. |last5=Webster |first5=G. |journal=Antonie van Leeuwenhoek|volume=100|issue=1|pages=83–98|s2cid=10224623}}</ref> ''Osedax'' is a monogeneric clade specialized in living on whale bones, although recent evidence shows them living on fish bones as well.<ref>{{cite journal|doi=10.1098/rsbl.2011.0202|title=Not whale-fall specialists, Osedax worms also consume fishbones|year=2011 |last1=Rouse |first1=G. W. |last2=Goffredi |first2=S. K. |last3=Johnson |first3=S. B. |last4=Vrijenhoek |first4=R. C.|journal=Biology Letters|volume=7|issue=5|pages=736–739|pmid=21490008|pmc=3169056}}</ref>

One probable relationship between the four clades is shown in the cladogram below. The position of ''Osedax'' is weakly supported.<ref name="ReferenceA"/>
{{clade
⚫
|label1=Siboglinidae
|1={{clade
|1=clade Frenulata
|2={{clade
|1=''Osedax''
|2={{clade
|1=''Sclerolinum'' (clade Monilifera)
|2=clade Vestimentifera
}}
}}
}}
}}


==Vestimentiferans==
==Vestimentiferans==
Like other tube worms, vestimentiferans are [[Benthic zone|benthic]] marine creatures. ''[[Riftia pachyptila]]'', a vestimentiferan, is known only from the [[hydrothermal vent]] systems.<ref name="ReferenceA"/>
Like other tube worms, vestimentiferans are marine and [[wiktionary:benthic|benthic]]. ''[[Riftia pachyptila]]'', a vestimentiferan, is known only from the [[hydrothermal vent]] systems.<ref name="ReferenceA"/> The vestimentiferans possess an anterior first body part called the obturaculum. Their main trunk of the body bears winglike extensions, the vestimentum, from which their name is derived. Also, unlike other siboglinids that never have a [[digestive tract]], they have one that they completely lose during [[metamorphosis]]. Their primary nutrition is derived from the sulphide-rich fluids emanating from the hydrothermal vents they live by. The sulphides 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. It is not completely understood how the worms instigate their relationship with the bacteria. One theory is that the very young worm has a vent on its body permitting the entry of the bacteria from the water.<ref name=Extra/> A more recent study of three species of tubeworms including ''Riftia pachyptila'' demonstrated that the bacteria actually infect juvenile worms through their skin. Their body is divided into four regions; the obturaculum, vestimentum, trunk, and [[opisthosome]].

===Anatomy of vestimentiferans===
[[File:Lamellibrachia satsuma.png|thumb|''[[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.<ref name="Miyamoto2013">{{cite journal|last1=Miyamoto|first1= Norio |last2=Shinozaki |first2= Ayuta|last3=Fujiwara |first3= Yoshihiro |date=2013-01-23|page=e55151 |title=Neuroanatomy of the Vestimentiferan Tubeworm Lamellibrachia satsuma Provides Insights into the Evolution of the Polychaete Nervous System |volume=8 |doi=10.1371/journal.pone.0055151 |journal= PLOS ONE|issue= 1 |pmid= 23372830 |pmc= 3553155 |url= https://www.researchgate.net/publication/235393504|doi-access= free }}</ref> 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.<ref name="Miyamoto2013"/> 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.<ref>{{Cite journal|last1=Nussbaumer|first1=Andrea D.|last2=Fisher|first2=Charles R.|last3=Bright|first3=Monika|date=2006-05-18|title=Horizontal endosymbiont transmission in hydrothermal vent tubeworms|journal=Nature|volume=441|issue=7091|pages=345–348|doi=10.1038/nature04793|issn=1476-4687|pmid=16710420|bibcode=2006Natur.441..345N|s2cid=18356960}}</ref> 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.<ref>{{Cite journal|last1=Reveillaud|first1=Julie|last2=Anderson|first2=Rika|last3=Reves-Sohn|first3=Sintra|last4=Cavanaugh|first4=Colleen|last5=Huber|first5=Julie A.|date=2018-01-27|title=Metagenomic investigation of vestimentiferan tubeworm endosymbionts from Mid-Cayman Rise reveals new insights into metabolism and diversity|journal=Microbiome|volume=6|issue=1|pages=19|doi=10.1186/s40168-018-0411-x|issn=2049-2618|pmc=5787263|pmid=29374496 |doi-access=free }}</ref> When the host dies, the bacteria are released and return to the free-living population in the seawater.<ref>{{Cite journal|last1=Klose|first1=Julia|last2=Polz|first2=Martin F.|last3=Wagner|first3=Michael|last4=Schimak|first4=Mario P.|last5=Gollner|first5=Sabine|last6=Bright|first6=Monika|date=2015-09-08|title=Endosymbionts escape dead hydrothermal vent tubeworms to enrich the free-living population|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=112|issue=36|pages=11300–11305|doi=10.1073/pnas.1501160112|issn=1091-6490|pmc=4568656|pmid=26283348|bibcode=2015PNAS..11211300K|doi-access=free}}</ref>

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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 [[Cold seep|hydrocarbon seep]] by roots located at the basal portion of their bodies.<ref>{{cite journal|doi=10.1007/s002270050233 |last1=Halanych |first1=K. |last2=Maas |first2=P. |last3=Hoeh |first3=W. |last4=Hashimoto |first4=J. |last5=Desbruyeres |first5=D. |last6=Lutz |first6=R. |last7=Vrijenhoek |first7=R. |title=Molecular systematics of vestimentiferan tubeworms from hydrothermal vents and cold-water seeps|year=1997|journal=Marine Biology|volume=130|issue=2|pages=141–149|s2cid=13217485 }}</ref> 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&nbsp;m.{{Citation needed|date=May 2023}}

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.<ref>{{cite journal|pmid=10441078 |last1=Julian |first1=D. |last2=Gaill |first2=F. |last3=Wood |first3=E. |last4=Arp |first4=A. |last5=Fisher |first5=C. |title=Roots as a site of hydrogen sulphide uptake in the hydrocarbon seep vestimentiferan ''Lamellibrachia'' sp |url=http://jeb.biologists.org/content/202/17/2245.short |year=1999|volume=202|issue=Pt 17|pages=2245–57|journal=The Journal of Experimental Biology|doi=10.1242/jeb.202.17.2245 }}</ref> 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.
⚫
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.<ref>{{cite journal|doi=10.1007/s002270050233|author=Halanych, K., Maas, P., Hoeh, W., Hashimoto, J., Desbruyeres, D., Lutz, R., and Vrijenhoek, R.|title=Molecular systematics of vestimentiferan tubeworms from hydrothermal vents and cold-water seeps|year=1997|journal=Marine Biology|volume=130|issue=2|pages=141–149}}</ref> 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. It is unsure how long the roots of the tube worms can grow to, but root have been recovered longer than 30&nbsp;m in length. 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.<ref>{{cite journal|pmid=10441078|author= Julian, D., Gaill, F., Wood, E., Arp, A., and Fisher, C|title=Roots as a site of hydrogen sulphide uptake in the hydrocarbon seep vestimentiferan Lamellibrachia sp|year=1999|volume=202|issue=Pt 17|pages=2245–57|journal=The Journal of Experimental Biology}}</ref> 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 tubeworms 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. The Vestimentiferan tubeworms build the external chitin structure themselves by secreting chitin from specialized glands located in their body walls.


==Genera in the family Siboglinidae==
==Genera==
* ''[[Osedax]]''
* ''[[Osedax]]''
* Frenulata<ref name=Rouse/>
* Clade ''[[Frenulata]]''<ref name=Rouse/>
** ''[[Birsteinia]]''
** ''[[Birsteinia]]''
** ''[[Bobmarleya]]''<ref>Ana Hilário and Marina R. Cunha, On some frenulate species (Annelida: Polychaeta: Siboglinidae) from mud volcanoes in the Gulf of Cadiz (NE Atlantic)</ref>
** ''[[Bobmarleya]]''<ref>{{cite journal |first1=A. |last1=Hilário |first2=M. R. |last2=Cunha |title=On some frenulate species (Annelida: Polychaeta: Siboglinidae) from mud volcanoes in the Gulf of Cadiz (NE Atlantic) |journal=Scientia Marina |volume=72 |issue=2 |year=2008 |pages=361–371 |doi=10.3989/scimar.2008.72n2361|doi-access=free }}</ref>
** ''[[Choanophorus]]''
** ''[[Choanophorus]]''
** ''[[Crassibrachia]]''
** ''[[Crassibrachia]]''
Line 47: Line 70:
** ''[[Diplobrachia]]''
** ''[[Diplobrachia]]''
** ''[[Galathealinum]]''
** ''[[Galathealinum]]''
** ''[[Heptobrachia]]''
** ''[[Heptabrachia]]''
** ''[[Lamellisabella]]''
** ''[[Lamellisabella]]''
** ''[[Nereilinum]]''
** ''[[Nereilinum]]''
** ''[[Oligobrachia]]''
** ''[[Oligobrachia]]''
** ''[[Paraescarpia]]''<ref>{{cite journal | author = Southward ''et al.'' | year = 2002 | title = Vestimentiferans (Pogonophora) in the PaciŽ c and Indian Oceans: a new genus from Lihir Island (Papua New Guinea) and the Java Trench, with the Ž rst report of Arcovestia ivanovi from the North Fiji Basin | url = | journal = J. Nat. Hist. | volume = 36 | issue = | pages = 1179–1197 | doi=10.1080/00222930110040402}}</ref>
** ''[[Paraescarpia]]''<ref>{{cite journal | last1 = Southward | first1 = E. C. | first2 = A. | last2 = Schulze |first3 = V. | last3 = Tunnicliffe | year = 2002 | title = 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 = Journal of Natural History | volume = 36 | issue = 10 | pages = 1179–1197 | doi=10.1080/00222930110040402| s2cid = 86076917 }}</ref>
** ''[[Polybrachia]]''
** ''[[Polybrachia]]''
** ''[[Siboglinoides]]''
** ''[[Siboglinoides]]''
Line 58: Line 81:
** ''[[Spirobrachia]]''
** ''[[Spirobrachia]]''
** ''[[Unibrachium]]''
** ''[[Unibrachium]]''
** ''[[Volvobrachia]]''<ref>{{cite journal | last=Smirnov |first=R. V. | year=2000 | title=A redescription of Spirobrachia leospira Gureeva (Pogonophora), with the erection of a new genus and a revision of the Spirobrachiidae | journal=Ophelia | volume=53 | issue=2 | pages=151–158 | doi=10.1080/00785236.2000.10409445|s2cid=85319259 }}</ref>
** ''[[Volvobrachia]]''{{citation needed|date=November 2012}}
** ''[[Zenkevitchiana]]''
** ''[[Zenkevitchiana]]''
* Clade Monilifera<ref>{{Cite journal |last=Halanych |first=Kenneth M. |date=2005 |title=Molecular phylogeny of siboglinid annelids (a.k.a. pogonophorans): a review |journal=Hydrobiologia |volume=535-536 |issue=1 |pages=297–307 |doi=10.1007/s10750-004-1437-6 |s2cid=16022909 |issn=0018-8158}}</ref>
⚫
* ''[[Sclerolinum]]''<ref name=Rouse/>
* Vestimentifera<ref name=Rouse/>
** ''[[Sclerolinum]]''<ref name=Rouse/>
⚫
* Clade ''[[Vestimentifera]]''<ref name=Rouse/>
** ''[[Alaysia]]''
** ''[[Alaysia]]''
** ''[[Arcovesia]]''
** ''[[Arcovesia]]''
Line 71: Line 95:
** ''[[Tevnia]]''
** ''[[Tevnia]]''


== References ==
==References==
{{Reflist}}
⚫
{{Commons category|Pogonophora}}
⚫
{{Wikispecies|Siboglinidae}}
{{reflist}}


==External links==
{{Animalia}}
⚫
*{{Commons category-inline|Siboglinidae}}
⚫
*{{Wikispecies-inline|Siboglinidae}}


{{Taxonbar|from=Q809618}}
{{Authority control}}
{{Authority control}}


[[Category:Sabellida]]
[[Category:Polychaetes]]
[[Category:Chemosynthetic symbiosis]]
[[Category:Annelid families]]

Revision as of 14:08, 29 April 2024

Siboglinidae
Temporal range: 189.6–Recent Ma
Riftia pachyptila
Scientific classification Edit this classification
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.

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] Molecular work aligning five genes has identified four distinct clades within Siboglinidae.[7][8][9] The clades are Vestimentifera, Sclerolinum, Frenulata, and Osedax.[8] Vestimentiferans live in vent and seep habitats.[8] 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.[10] Sclerolinum is a monogeneric clade (which may be called Monilifera) living on organic-rich remains.[5] Frenulates live in organic-rich sediment habitats.[11] Osedax is a monogeneric clade specialized in living on whale bones, although recent evidence shows them living on fish bones as well.[12]

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

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.[13] 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.[13] 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.[14] 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.[15] When the host dies, the bacteria are released and return to the free-living population in the seawater.[16]

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.[17] 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.[18] 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

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. ^ a b c d 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. ^ a b 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. ^ a b c d e f 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. ^ 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.
  8. ^ a b c 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.
  9. ^ 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.
  10. ^ 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
  11. ^ 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.
  12. ^ 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.
  13. ^ a b 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.
  14. ^ 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.
  15. ^ 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.
  16. ^ 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.
  17. ^ 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.
  18. ^ 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.
  19. ^ 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.
  20. ^ 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.
  21. ^ 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.
  22. ^ 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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