# Osedax

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***Osedax*** is a [genus](/source/Genus) of [siboglinid](/source/Siboglinid) [polychaetes](/source/Polychaete), commonly called **snot worms** or **bone-eating worms**. *Osedax* is [Latin](/source/Latin) for 'bone devourer'[1], derived from the worms' unique ecological niche of bone-boring. *Osedax* settle on a bone, then secrete an acid through specialized root tissues to dissolve the [bone's external layers](/source/Bone#Cortex) in order to access the [lipids](/source/Lipid) within.[1][2][3] *Osedax* act as [ecosystem engineers](/source/Ecosystem_engineer), enhancing the biodiversity of bones they inhabit by increasing their structural complexity, allowing [microfauna](/source/Microfauna) to inhabit otherwise inaccessible regions of the internal bone.[3]

Scientists from the [Monterey Bay Aquarium Research Institute](/source/Monterey_Bay_Aquarium_Research_Institute) using the submarine ROV *Tiburon* first discovered the genus in [Monterey Bay](/source/Monterey_Bay), [California](/source/California), in February 2002. The worms were found living on the bones of a decaying [gray whale](/source/Gray_whale) in the [Monterey Canyon](/source/Monterey_Canyon), at a depth of 2,893 m (9,491 ft).[1]

## Anatomy and physiology

Like other [siboglinids](/source/Siboglinid), *Osedax* lacks a mouth, gut, or anus, and instead depends on colonies of [endosymbiont](/source/Endosymbiont) microbes housed inside a [trophosome](/source/Trophosome) for nutrition. Unlike other siboglinids, however, this trophosome takes the form of a vascularized root system which penetrates bone.[1][4] These microbes, of the order *[Oceanospirillales](/source/Oceanospirillales),* produce enzymes which [hydrolyze](/source/Hydrolysis) [collagen](/source/Collagen) from bones, yielding nutrition to the worms.[4][1][5][6][7] *Osedax* exhibit very drastic sexual dimorphism, with females being >20,000 times larger than males.[8] Males are [paedomorphic](/source/Neoteny) and microscopic, inhabiting a section of the females' [trunk](/source/Siboglinidae#Anatomy) where they produce sperm from yolk reserves.[1][8]

### Digestive system

*Osedax* rely on [symbiotic](/source/Symbiosis) species of bacteria that aid in the digestion of whale proteins and lipids and release nutrients that the worms can absorb.[1] *Osedax* have colorful feathery plumes that also act as [gills](/source/Gill) and unusual root-like structures that absorb nutrients. The *Osedax* secrete acid (rather than rely on teeth) to bore into bone to access the nutrients[1][2]. High concentrations of [carbonic anhydrase](/source/Carbonic_anhydrase) are found in the roots of *Osedax.* This serves as evidence of a common bioerosion mechanism in which secreted acid is produced by [aerobic respiration](/source/Aerobic_respiration). This process works with a demineralization mechanism in which oxygen is carried from seawater to the roots and HCO is secreted into the seawater.[2]

The epidermis also plays key roles in bone deterioration and nutrient uptake. This process of bone deterioration occurs through a symbiotic relationship with an [endosymbiotic](/source/Endosymbiont) bacteria.[9] The cells in the epidermis of the *Osedax* root region are responsible for the secretion of digestive enzymes. The epidermis also has an expanded microvillus border which increases the surface area.[9]

Through the use of [X-ray CT](/source/X-ray_crystallography) technology, scans showed that borings made by *Osedax mucofloris* were hemi-ellipsoidal in shape. Boring depths varied depending on which bone was colonized by the *O. mucofloris*. Deeper borings were found in radius bone compared to the ulna and vertebrae.[10]

*Osedax* roots are covered by a mucus sheath that helps protect the worm's trunk. Some studies support the theory that this sheath plays a role in dissolving the bone. This sheath could also play an important role in reducing the damage to *Osedax* skin by absorbing harmful acid. Another potential function of the mucus sheath is that it could inhibit the breakdown of the worm's bone matrix. This is significant because the bone matrix is integral in maintaining the worm's position while in direct contact with a bone.[2]

### Symbiosis

Symbionts are the primary providers of nutrition for *Osedax.*[1] However, these symbionts also possess [genes](/source/Gene), [secretion systems](/source/Bacterial_secretion_system), and [toxins](/source/Toxin#Biological) that disrupt the *Osedax* membrane and facilitate recurrent infections of adult *Osedax* through the root tips.[1][11] There is ongoing debate in the literature over whether the [symbiosis](/source/Symbiosis) in *Osedax* roots is [commensal](/source/Commensalism) or [mutualistic](/source/Mutualism_(biology)).[12][11] The symbiotic relationship between *Osedax* and its accompanying bacteria may be transferred either via [vertical](/source/Vertical_transmission) or [horizontal transmission](/source/Horizontal_transmission).[11]

*Osedax* species use collagen, which is the primary organic component in bone.[7] Collagen is degraded using a family of [endopeptidases](/source/Endopeptidase) called [matrix metalloproteinases (MMPs)](/source/Matrix_metalloproteinase), which facilitates nutrient absorption by the *Osedax*.[7] The roots of the *Osedax* express high amounts of [V-ATPase](/source/V-ATPase) and carbonic anhydrase [enzymes](/source/Enzyme), which allows the *Osedax* to dissolve and absorb collagen and [lipids](/source/Lipid).[7] Once dissolved, the nutrients are either used by the *Osedax*, or transported to the symbionts for further catabolism.[11][7]

As the [endosymbionts](/source/Endosymbiont) lack secreted M9 peptidase, they rely on the *Osedax* worm to source extracellular collagen.[7] The symbionts in the *[Oceanospirillales](/source/Oceanospirillales)* order have then been observed to further process the collagen using collagenolytic enzymes.[11][6]

[Sequencing](/source/DNA_sequencing) of the *Osedax* worm [genome](/source/Genome) has suggested an evolved dependency on its endosymbionts.[7] This is revealed by genomic streamlining, where increased functional groups were observed despite the loss of some gene families.[7] Six incomplete pathways were discovered in the *Osedax* worm genome which were supplemented by the endosymbionts.[7] In particular, the *Osedax* worm lacks specific gene families involved in bone lipid and [carbohydrate metabolism](/source/Carbohydrate_metabolism).[7] This function is complemented by the *Oceanospirillales* symbionts, which utilize the [glyoxylate cycle](/source/Glyoxylate_cycle) to catabolize nutrients from whale bones and convert [fatty acids](/source/Fatty_acid) into [carbohydrates](/source/Carbohydrate).[7] The *Osedax* are then able to take up and store the end products as [glycogen](/source/Glycogen).[7] [Bacteriocytes](/source/Bacteriocyte) are present in the *Osedax* lower trunk subepidermal connective tissue,[7] and there are additional genes in the bacteriocytes that encode [amino acids](/source/Amino_acid) and [glucose](/source/Glucose) and aid in digestion and absorption of [proteins](/source/Protein) into the roots.[13]

### Endosymbionts

The *[Oceanospirillales](/source/Oceanospirillales)* symbionts are found in the specialized roots[12] of all *Osedax* species,[14][11] and play a major role in accelerating the degradation process of bones, as well as facilitating nutrient uptake for the *Osedax*.[12][7] *Oceanospirillales* are known for their ability to degrade complex organic compounds.[1][6]

*[Campylobacterales](/source/Campylobacterales)* are abundant along the trunk of the *Osedax* according to a 2023 study.[12] Different [genera](/source/Genus) in this order are found in *Osedax* at different points during the whale's degradation:

1. Members of the *[Arcobacter](/source/Arcobacter)* genus are the primary early colonizers (<24 months).[12]
1. *[Sulfurospirillum](/source/Sulfurospirillum)* genus members colonize at ~50 months, during the transitional stages of organic carbon breakdown.[12]
1. The *[Sulfurimonas](/source/Sulfurimonas)* genus dominates at >140 months, and are key players in its symbiosis with the *Osedax* host.[12]

The *Sulfurimonas* genus in particular protects the *Osedax* worms from potentially harmful by-products produced at >140 months of the whale fall degradation.[12] The *Sulfurimonas* bacteria house the type II and IV [sulfide:quinone oxidoreductase](/source/Sulfide:quinone_reductase) genes that encode enzymes to oxidize and assimilate sulfide.[12] These reactions prevent the host from absorbing toxic by-products across the [epithelial barrier](/source/Epithelium).

## Sexual dimorphism and life cycle

Osedax males are notably smaller than their female counterparts. Between 50 and 100 microscopic dwarf males live inside the tube surrounding a single female and never develop past the larval stage; they produce sperm from yolk reserves.[8] Male dwarfism prevents competition with female *Osedax* worms for food and space.[8] Conditions that favour dwarfism in male Osedax are:

1. Eliminates competition between male and female *Osedax* as resources are limited,[15]
1. Sessile lifestyle: attach to and rely on females for food,[15]
1. Decreases difficulty in finding a mate.[15]

Interestingly, *[Osedax priapus](/source/Osedax_priapus)* lack the frequently observed male dwarfism. Males live freely and compete for space and food with females. Being larger, they produce and carry more sperm. However, sexual size dimorphism is still observed in *O. priapus*: most males are one-third the volume of females.[8]

Annelid sex is typically determined by genetic factors,[8] however models of environmental sex determination have been proposed for *Osedax*, in which larvae that settle on bones mature into females, while larvae that settle on female *Osedax* do not fully develop and mature into males.[1] *[Osedax japonicus](/source/Osedax_japonicus)* in particular has showcased an environmental form of sex determination.[8]

### Life cycle

Female *Osedax* worms have been observed [spawning](/source/Spawn_(biology)) both in the wild and in laboratory aquaria.[8] *[Osedax rubiplumus](/source/Osedax_rubiplumus)* can spawn hundreds of [oocytes](/source/Oocyte) at a time. They are already fertilized when they are released from the female worm. The worms' endosymbionts, species of bacteria in the order [Oceanospirillales](/source/Oceanospirillales), were not observed in the spawned oocytes, which suggests that they are acquired after the worms settle on the bones.[8] *Osedax* appears to be highly fecund and reproduces continuously.

- Mature female *Osedax* worms spawn eggs into the mucus attached to their tubes, where the embryos develop for 3 days.
- Larvae then begin to swim in the water column. This is called the trochophore stage. The larvae settle on whale bones and begin crawling.
- During the trocophore stage, male *Osedax* settle on the tubes of the females, where they are metamorphosed into dwarf males, which can be inside or outside the female tube.
- 1 day after settling on bones, larvae use two pairs of chaetae to attach to the substrate. Juvenile worms begin to secrete mucus and develop two ventral palps on the dorsal side of the prostomium.
- 2 days after settling, the palps elongate and the heart starts to beat. The roots attach to the bones begin to digest.
- 4 days after settling, the trunk and ventral palps elongate, where symbiotic bacteria are detected in the root.
- 7 days after settlement, pinnules extend from the ventral palps.
- 10 days post settlement, the juvenile worms have 4 palps with pinnules, an oviduct, and a distinct root system.[8]

## Ecology

When initially discovered, some scientists proposed that *Osedax* was a whale-bone specialist[16], a notion later disproven by the discovery of *Osedax* on a variety of skeletal substrates including bone fragments[17], [teleost](/source/Teleost) bones[14][17], shark teeth[18][19], reptile[20][21] and bird skeletons[22][17], pinniped bones[17], experimentally placed cow bones[23], and even human remains[24]. Multiple species of *Osedax* can co-exist in close proximity, even on the same bone.[25][17]

*Osedax* and their borings enhance local [biodiversity](/source/Biodiversity) by dissolving the tough external [cortical bone](/source/Bone#Cortex) and tunneling through their internal layers, increasing structural complexity and providing new regions for [microfauna](/source/Microfauna) to colonize.[3] As such, *Osedax* are considered [ecosystem engineers](/source/Ecosystem_engineer). However, these borings compromise the structural integrity of the bones, causing them to collapse after extensive boring, rendering these new habitats temporary.[3] The role of *Osedax* in the degradation of marine vertebrate remains is important to marine vertebrate [taphonomy](/source/Taphonomy).

## Evolution

The oldest trace fossils on bones characteristic of *Osedax* are from a [plesiosaur](/source/Plesiosauria) humerus from the [Cambridge Greensand](/source/Cambridge_Greensand), England, likely [reworked](/source/Reworked_fossil) from late [Albian](/source/Albian) (c. 100 million years old) sediments and a rib and costal plate from a [sea turtle](/source/Sea_turtle) found in [Cenomanian](/source/Cenomanian) (100–93 million years ago) aged sediments of the [Chalk Group](/source/Chalk_Group), England.[20] Further material is known from the [Campanian](/source/Campanian) and [Maastrichtian](/source/Maastrichtian).[26] Following the extinction of almost all large marine reptiles at the end of the [Cretaceous](/source/Cretaceous), *Osedax* likely persisted on the bones of sea turtles, marine birds, and fish.[27]

## Species

Selected species:[28][17][29][30]

- *[Osedax antarcticus](/source/Osedax_antarcticus)* Glover, Wiklund & Dahlgren, 2013
- *Osedax bozoi* Berman, Hiley, Read & Rouse, 2024
- *Osedax braziliensis* Fujiwara, Jimi, Sumida, Kawato, Kitazato
- *Osedax bryani* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax byronbayensis* Georgieva, Wiklund, Ramos, Neal, Glasby & Gunton, 2023
- *Osedax craigmcclaini* Berman, Hiley, Read, Rouse, 2024
- *[Osedax crouchi](/source/Osedax_crouchi)* Amon, Wiklund, Dahlgren, Copley, Smith, Jamieson & Glover, 2014
- *[Osedax deceptionensis](/source/Osedax_deceptionensis)* Taboada, Cristobo, Avila, Wiklund & Glover, 2013
- *Osedax docricketts* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax estcourti* Berman, Hiley, Read & Rouse, 2024
- *Osedax fenrisi* Eilertsen, Dahlgren & Rapp, 2020
- *[Osedax frankpressi](/source/Osedax_frankpressi)* Rouse, Goffredi & Vrijenhoek, 2004
- *Osedax jabba* Rouse, Goffredi, Johnson & Vrijenhoek
- *[Osedax japonicus](/source/Osedax_japonicus)* Fujikura, [Fujiwara](/source/Yoshihiro_Fujiwara) & [Kawato](/source/Masaru_Kawato), 2006
- *Osedax knutei* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax lehmani* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax lonnyi* Rouse, Goffredi, Johnson & Vrijenhoek
- *[Osedax mucofloris](/source/Osedax_mucofloris)* Glover, Kallstrom, Smith & Dahlgren, 2005
- *Osedax nataliae* Gularte, Sumida, Bergamo & Rouse, 2024
- *[Osedax nordenskjoeldi](/source/Osedax_nordenskjoeldi)* Amon, Wiklund, Dahlgren, Copley, Smith, Jamieson & Glover, 2014
- *[Osedax priapus](/source/Osedax_priapus)* Rouse et al., 2014
- *Osedax packardorum* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax randyi* Rouse, Goffredi, Johnson & Vrijenhoek
- *[Osedax rogersi](/source/Osedax_rogersi)* Amon, Wiklund, Dahlgren, Copley, Smith, Jamieson & Glover, 2014
- *[Osedax roseus](/source/Osedax_roseus)* Rouse, Worsaae, Johnson, Jones & Vrijenhoek, 2008
- *[Osedax rubiplumus](/source/Osedax_rubiplumus)* Rouse, Goffredi & Vrijenhoek, 2004
- *Osedax ryderi* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax sigridae* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax talkovici* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax tiburon* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax traceyae* Berman, Hiley, Read & Rouse, 2024
- *Osedax ventana* Rouse, Goffredi, Johnson & Vrijenhoek
- *Osedax waadjum* Georgieva, Wiklund, Ramos, Neal, Glasby & Gunton, 2023
- *Osedax westernflyer* Rouse, Goffredi, Johnson & Vrijenhoek

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1. [WoRMS, Genus Osedax](http://www.marinespecies.org/aphia.php?p=taxdetails&id=265008)

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## External links

- [Press release describing discovery of *Osedax*](https://web.archive.org/web/20040805034842/http://www.mbari.org/news/news_releases/2004/whalefall.html)
- [BBC website](https://news.bbc.co.uk/2/hi/science/nature/4354286.stm) – link to story about discovery of *Osedax* worms in the North Sea
- [A Motley Collection of Boneworms – Monterey Bay Aquarium Research Institute](https://web.archive.org/web/20091114064117/http://www.mbari.org/news/news_releases/2009/osedax-spp/osedax-spp-release.html)
- [Discovered in the deep: the worm that eats bones](https://www.theguardian.com/environment/2022/aug/22/discovered-in-the-deep-the-worm-that-eats-bones-osedax) – The Guardian

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Adapted from the Wikipedia article [Osedax](https://en.wikipedia.org/wiki/Osedax) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Osedax?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.