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Oxyrrhis

Oxyrrhis /ˌɑkˈsɪɹɪs/ is a genus of heterotrophic dinoflagellate, the only genus in the family Oxyrrhinaceae. It inhabits a range of marine environments worldwide and is important in the food web dynamics of these ecosystems. It has the potential to be considered a model organism for the study of other protists. Oxyrrhis is an early-branching lineage and has long been described in literature as a monospecific genus, containing only Oxyrrhis marina. Some recent molecular phylogenetic studies argue that Oxyrrhis comprises O. marina and O. maritima as distinct species, while other publications state that the two are genetically diverse lineages of the same species. The genus has previously been suggested to contain O. parasitica as a separate species, however the current consensus appears to exclude this, with Oxyrrhis being monospecific and containing O. marina and O. maritima as separate lineages of the type species. The genus is characterised by its elongated body which is anteriorly prolonged to a point, its complex flagellar apparatuses which attach to the ventral side of the cell, and the unique features of its nucleus.

Etymology
The name Oxyrrhis is derived from the Greek oxys, meaning 'sharp' and rhis, meaning 'nose'. This indicates the anterior extension of the body. == Type species ==
Type species
The type species for this genus is Oxyrrhis marina. == History of knowledge ==
History of knowledge
Oxyrrhis was first described by Félix Dujardin in 1841, having been discovered in a salt marsh habitat in Belgium. The amount of knowledge relating to O. marina is rapidly increasing, with the number of publications having greatly risen in the 1990s. Additionally, Oxyrrhis is a food source for several planktivores, so there is some data relating to predation rates upon it. Between 1938 and 2009, 144 papers were published that featured Oxyrrhis as a keyword. In a survey of 36 papers from 1990-2011 that mention O. marina in the title, approximately 55% were aut- or synecologically based, roughly 40% examined an aspect related to evolutionary or genetic biology, and 5% were associated with distributional patterns. However, only around 5-10% of these papers showed any evidence of rigorous cross-disciplinary evaluation. == Habitat and ecology ==
Habitat and ecology
Oxyrrhis is widely regarded as having global distribution, but there are limited studies of its geographic range. Most published data describe the range of Oxyrrhis as areas of the Atlantic and Pacific coasts of the USA, the Gulf of Mexico, the Atlantic coasts of Europe, the Mediterranean and Baltic Seas, the Persian Gulf, the Indian Ocean, and the western Pacific. Its presence has been confirmed in the southern hemisphere, such as in or around Australia, South Africa, Brazil, and it is rare or absent in polar seas (northern Norway, Iceland), however there have been small sample sizes in these polar regions. Oxyrrhis is rare in open water but has been found to inhabit the coastal waters of some remote islands including Hawaii and the Azores. The genus shows both widespread distributions and endemicity through its various clades. It has been discovered that one clade of O. marina has widespread distribution, covering both coasts of North America and the eastern Mediterranean Sea, while the other clade is only found in the Baltic Sea and Red Sea. In terms of habitat, Oxyrrhis is common in many intertidal and coastal habitats. There is potential sampling bias when assessing its distribution, as virtually all samples have been from coastal areas, mainly intertidal pools, rather than the open ocean. Oxyrrhis is unlikely to be exclusively intertidal and is probably a small component of coastal and oceanic plankton communities, while there is anecdotal evidence that it occasionally grows in shallow embayments. Differences in the temperature and salinity ranges of O. marina and O. maritima suggest that the two lineages have different ecophysiology. O. maritima lives in tidal pools with a relatively high salinity and temperature, while O. marina lives in water columns which have a lower salinity and temperature. The highest temperature for positive growth of O. marina is likely to be around 32 °C and 32 °C or higher for O. maritima. O. marina only grows at a salinity >=4 ppt and the growth rate increases with salinity up to 50 ppt. O. maritima grows at a salinity of 2 ppt, and growth rate also increases up until 50 ppt. Oxyrrhis is highly important in marine communities, playing an essential role in pelagic food webs as they both consume phytoplankton in addition to ciliates, bacteria, and the eggs, early nauplii stages, and adults of some metazoans, amongst other prey items. They additionally act as prey for upper trophic levels, which include some metazoans, ciliates and other dinoflagellates. These consumption rates by Oxyrrhis can exceed 60% of the daily phytoplankton production in many of oceanic and coastal systems. By feeding on and being predated upon by a broad range of organisms, Oxyrrhis significantly affect food web structure, carbon cycles and energy flows within the marine planktonic community. == Description of the organism ==
Description of the organism
Morphology/anatomy Dujardin described Oxyrrhis as easily recognisable by its oblong, irregular, obliquely truncated shape, and by its flagelliform filaments. Oxyrrhis might possess feeding receptors such as contact chemoreceptors or lectin-like receptors to recognize carbohydrate moieties, which are cell surface components of their marine microalgal and bacterial prey, allowing for highly specific ingestion behaviours. These apparatuses are asymmetric and very complex, with each flagellum having a longitudinal and a transverse basal body, giving rise to eight structurally different components. Some aspects of Oxyrrhis nuclear and chromosomal organization are unique, distinguishing it from other genera. For example, rather than using an extranuclear spindle as typically seen in dinoflagellates, mitosis in Oxyrrhis is facilitated by the use of an intranuclear spindle. When triggered, haploid cells either fuse to form a motile diploid zygote or form a resting cyst, taking on the role of gametes in either case. Its genome is highly fragmented, like that of other dinoflagellates, but the genes are frequently arranged as tandem copies, similar to the repeating nature of the Plasmodium genome. The genome of Oxyrrhis appears to be structured as neighboring genes or gene fragments, and these are always the same. This means the cox1 and the cob-cox3 fusion are never found on the same genomic fragment. Neither cox1, cob-cox3, nor circularized mRNAs from cob-cox3 have been found to use canonical start or stop codons. Additionally, Oxyrrhis does not exhibit the extensive RNA editing that is characteristic of the dinoflagellates. == Practical importance ==
Practical importance
O. marina meets a range of criteria that combined, could deem it suitable as a 'model organism' for answering many different types of questions relating to other similar protists. This has given insight into how changing distributions of chemical cues in the environment can influence predator-prey interactions among related genera. During laboratory experiments, O. marina effectively controls populations of H. akashiwo through grazing. Calculated impacts on natural populations suggest that large-scale culturing of O. marina could be used to successfully manage the red tides that result from H. akashiwo blooms. Additionally, Oxyrrhis is used in fish food, to enhance larval survival in aquaculture and the tropical fish trade due to its ease of mass culturing and potentially high nutritional quality. The feeding of Oxyrrhis to larval fish in aquaculture could lead into future research on the role of protozoa as a food source for larval fish in natural waters. == List of species and lower taxonomic units ==
List of species and lower taxonomic units
Species: O. marina Diverse lineages within species: O. marina and O. maritima, each comprising two clades, giving four clades in total which contain many strains. ==References==
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