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Siphonophore

Siphonophores are cnidarian animals of the hydrozoan order Siphonophorae. According to the World Register of Marine Species, the order contains 194 species described thus far.

Anatomy and morphology
Colony characteristics Siphonophores are colonial hydrozoans that do not exhibit alternation of generations, but instead reproduce asexually through a budding process. Zooids are the multicellular units that build the colonies of a siphonophore. A single bud called the pro-bud initiates the growth of a colony by undergoing fission. Cystonects have a long stem with attached zooids. Moreover, siphonophores possess multiple types of zooids. Scientists have determined two possible evolutionary hypotheses for this observation: 1. As time progresses, zooid types have increased. In general, siphonophore colonies have a modular body plan, with many different zooids making up the overall structure. These types can include: feeding gastrozooids, movement zooids, and sensory zooids. :Specifically, feeding zooids in siphonophores have undergone many unique adaptations to service the deep. Gastrozooids are uniquely specialized organisms that have feeding polyps (similar to a mouth) along with a long tentacle with side branches which is used to capture the prey. This adaptation is unique to zooids living in siphonophore colonies. Nectophores :Nectophores are medusae that assist in the propulsion and movement of some siphonophores in water. They are characteristic in physonectae and calycophores. The nectophores of these organisms are located in the nectosome where they can coordinate the swimming of colonies. They are gas-filled floats that are located at the anterior end of the colonies in these species. ==Distribution and habitat==
Distribution and habitat
Currently, the World Register of Marine Species (WoRMS) identifies 175 species of siphonophores. They can differ greatly in terms of size and shape, which largely reflects the environment that they inhabit. Siphonophores are most often pelagic organisms, yet level species are benthic. Smaller, warm water siphonophores typically live in the epipelagic zone and use their tentacles to capture zooplankton and copepods. Larger siphonophores live in deeper waters, as they are generally longer and more fragile and must avoid strong currents. They mostly feed on larger prey, such as small fishes or crustaceans. The majority of siphonophores live in the deep sea and can be found in all of the oceans. Siphonophore species rarely only inhabit one location. Some species, however, can be confined to a specific range of depths or an area of the ocean. ==Behavior==
Behavior
Movement Siphonophores use a method of locomotion similar to jet propulsion. A siphonophore is a complex aggregate colony made up of many nectophores, which are clonal individuals that form by budding and are genetically identical. Depending on where each individual nectophore is positioned within the siphonophore, their function differs. The velum becomes smaller and more circular during times of forward propulsion compared to a large velum that is seen during refill periods. Their diets consist of a variety of copepods, other small crustaceans, cnidarians, ctenophores, and small fish. Some siphonophores, such as Praya dubia, have been observed to feed on other species in the same order. Generally, the diets of strong swimming siphonophores consist of smaller prey, and the diets of weak swimming siphonophores consist of larger prey. Research has shown that specific diets vary even between different individuals of the same species depending on their particular environment. A majority of siphonophores have gastrozooids that have a characteristic tentacle attached to the base of the zooid. This structural feature functions in assisting the organisms in catching prey. The nematocysts then shoot millions The gelatinous body plan allows for flexibility when catching prey, but the gelatinous adaptations are based on habitat. They swim around waiting for their long tentacles to encounter prey. In addition, siphonophores in a group denoted Erenna have the ability to generate bioluminescence and red fluorescence while its tentilla twitches in a way to mimic motions of small crustaceans and copepods. These actions entice the prey to move closer to the siphonophore, allowing it to trap and digest it. Predators of Siphonophores include narcomedusae, gastropods, other siphonophores, and large fish such as Mola mola. This species then proceeds to undergo "blanching," and emit a startlingly bright light. Nanomia, among other siphonophore species, can move both forward and backwards to escape a stimulus from the opposite side, using their nectophores for propulsion. While there are few observations of defensive behavior in situ, it is argued that these strategies help siphonophores evade predators. There is limited research on the mechanistic release of eudoxid fragments for reproduction, and studies are determining whether to consider them as clustered communities or individuals. Recent research has identified eudoxid tissue remodeling before release by a specified muscle, as well as a dispersal mechanism that temporarily alters siphonophore buoyancy. Bioluminescence Nearly all siphonophores have bioluminescent capabilities. Since these organisms are extremely fragile, they are rarely observed alive. Bioluminescence in siphonophores has been thought to have evolved as a defense mechanism. Siphonophores of the deep-sea genus Erenna (found at depths between ) are thought to use their bioluminescent capability for offense too, as a lure to attract fish. This genus is one of the few to prey on fish rather than crustaceans. The bioluminescent organs, called tentilla, on these non-visual individuals emit red fluorescence along with a rhythmic flicking pattern, which attracts prey as it resembles smaller organisms such as zooplankton and copepods. Thus, it has been concluded that they use luminescence as a lure to attract prey. Some research indicates that deep-sea organisms can not detect long wavelengths, and red light has a long wavelength of 680 nm. If this is the case, then fish are not lured by Erenna, and there must be another explanation. However, the deep-sea remains largely unexplored and red light sensitivity in fish such as Cyclothone and the deep myctophid fish should not be discarded. Bioluminescent lures are found in many different species of siphonophores, and are used for a variety of reasons. Species such as Agalma okeni, Athorybia rosacea, Athorybia lucida, and Lychnafalma utricularia use their lures as a mimicry device to attract prey. A. rosacea mimic fish larvae, A. lucida are thought to mimic larvacean houses, and L. utricularia mimic hydromedusa. The species Resomia ornicephala uses their green and blue fluorescing tentilla to attract krill, helping them to outcompete other organisms that are hunting for the same prey. Erenna sirena uses bioluminescent lures surrounded by red fluorescence to attract prey and possibly mimic a fish from the Cyclothone genus. Their prey is lured in through a unique flicking behavior associated with the tentilla. When young, the tentilla of organisms in the Erenna genus contain only bioluminescent tissue, but, as the organism ages, red fluorescent material is also present in these tissues. ==Taxonomy==
Taxonomy
(Physalia physalis) History Carl Linnaeus described the first siphonophore, the Portuguese man o' war, in 1758. Another notable recent discovery occurred in early 2025, when a team led by researchers at Yale University, the University of New South Wales, and Griffith University determined that the Portuguese man o'war is actually at least four distinct species. The discovery has implications for scientist's understanding of open-ocean biodiversity, and encourages further research into understanding what caused and maintained this genetic variation. Modern taxonomy Siphonophores are classified into the phylum Cnidaria and the class Hydrozoa. The phylogenetic relationships of siphonophores have been of great interest due to the high variability of the organization of their polyp colonies and medusae. and floats (pneumatophores). The subgroups consisted of Cystonectae, Physonectae, and Calycorphores. Cystonectae had pneumatophores, Calycophores had nectophores, and Physonectae had both. Eukaryotic nuclear small subunit ribosomal gene 18S, eukaryotic mitochondrial large subunit ribosomal gene 16S, and transcriptome analyses further support the phylogenetic division of Siphonophorae into two main clades: Cystonectae and Codonophora. Suborders within Codonophora include Physonectae (consisting of the clades Calycophorae and Euphysonectae), Pyrostephidae, and Apolemiidae. • Suborder CalycophoraeAbylidae Agassiz, 1862 • Clausophyidae Totton, 1965 • Diphyidae Quoy & Gaimard, 1827 • Hippopodiidae Kölliker, 1853 • Prayidae Kölliker, 1853 • Sphaeronectidae Huxley, 1859 • Tottonophyidae Pugh, Dunn & Haddock, 2018 • Suborder CystonectaePhysaliidae Brandt, 1835 • Rhizophysidae Brandt, 1835 • Suborder PhysonectaeAgalmatidae Brandt, 1834 • Apolemiidae Huxley, 1859 • Cordagalmatidae Pugh, 2016 • Erennidae Pugh, 2001 • Forskaliidae Haeckel, 1888 • Physophoridae Eschscholtz, 1829 • Pyrostephidae Moser, 1925 • Resomiidae Pugh, 2006 • Rhodaliidae Haeckel, 1888 • Stephanomiidae Huxley, 1859 == Research challenges ==
Research challenges
For siphonophores, research has been limited. The organisms are gelatinous and fragile, making it difficult for researchers to obtain for sampling. Many siphonophores reside in the mesopelagic and bathypelagic zones, where many remotely operated vehicles (ROVs) and submersibles search to observe and sample these creatures. However, most colonies are destroyed during collections and because of this, there is a lack of preserved specimens for study. Most species of siphonophores are sampled through in situ observations, contributing to the gaps in knowledge about siphonophores and underrepresentation in species records because of sampling bias. Molecular phylogenetics has helped address some of the inequalities by using nuclear and mitochondrial gene sequences. Most of these scientific studies have provided more insight into the evolution of specialized zooids within colonies. Additionally, genomic analyses have used transcriptome datasets to improve phylogenetic resolution, allowing scientists and researchers to reconstruct patterns in evolutionary traits like the organization of colonies and the tentilla feeding structure. The manefish has been directly observed to associate with the Bathyphysa conifera, gaining shelter and access to food leftovers from the colony. ==References==
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