Siphonophores (from Ancient Greek σίφων (siphōn), meaning "tube" and -φόρος (-phóros), meaning "bearing") are cnidarian animals of the hydrozoan order Siphonophorae. According to the World Register of Marine Species, the order contains 194 species described thus far.
Siphonophores are highly polymorphic and complex organisms, which blur the line between individual organisms ("regular" animals, built from organs and cells) and colonial organisms (e.g., corals, where each polyp can in principle live on its own). Each specimen is composed of medusoid and polypoid zooids that are morphologically and functionally specialized. Zooids are multicellular units that develop from a single fertilized egg and combine to create functional colonies able to reproduce, digest, float, maintain body positioning, and use jet propulsion to move. Most colonies are long, thin, transparent floaters living in the pelagic zone.
Like other hydrozoans, some siphonophores emit light to attract and attack prey. While many sea animals produce blue and green bioluminescence, a siphonophore, Erenna sirena, was only the second life form found to produce a red light (the first being the scaleless dragonfish Chirostomias pliopterus).
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. Each zooid is produced to be genetically identical, however, mutations can alter their functions and increase diversity of the zooids within the colony. Siphonophores are unique in that the pro-bud initiates the production of diverse zooids with specific functions. The functions and organizations of the zooids in colonies vary widely among the different species, however the majority of colonies are bilaterally arranged with dorsal and ventral sides to the central stem (the stem is the vertical branch in the center of the colony to which the zooids attach). Zooids typically have special functions, and thus assume specific spatial patterns along the stem.
General morphology Siphonophores typically exhibit one of three standard body plans matching the suborders: Cystonectae, Physonectae, and Calycophorae. Cystonects have a long stem with attached zooids. Each group of zooids contains a gastrozooid. The gastrozooid consisting of a tentacle used for capturing and digesting food. The groups also have gonophores that are specialized for reproduction. They use a pneumatophore (a gas filled float), on their anterior end to drift at the surface of the water or stay afloat in the deep sea. Physonects have a pneumatophore and nectosome, which uses the nectophores for jet propulsion, in which the nectophores pump water backwards in order to move forward. Calycophorans differ from cystonects and physonects in that they have two nectophores and no pneumatophore. Instead, they typically possess oil filled glands that help with buoyancy.Jellyfish: A Natural History
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; and 2. the last common ancestor had many types of zooids and the diversity seen today is due to loss of zooid types. Research shows no evidence supporting the first hypothesis, and has seen some evidence in support of the second.
Zooids A zooid is a single part of an organism that makes up the greater whole. 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. The nectophores have also been observed in working in conjunction with reproductive structures in order to provide propulsion during colony detachment.
Bracts Bracts are zooids that are unique to the siphonophorae order. They function in protection and maintaining a neutral buoyancy. However, bracts are not present in all species of siphonophore.
Gastrozooids Gastrozooids are polyps that have evolved a function to assist in the feeding of siphonophores.
Palpons Palpons are modified gastrozooids that function in digestion by regulating the circulation of gastrovascular fluids.
Gonophores Gonophores are zooids that are involved in the reproductive processes of the siphonophores.
Pneumatophores The presence of pneumatophores characterizes the subgroups Cystonectae and Physonectae. They are gas-filled floats that are located at the anterior end of the colonies in these species. They function to help the colonies maintain their orientation in water. In the Cystonectae subgroup, the pneumatophores have an additional function of assisting with flotation of the organisms. The siphonophores exhibiting the feature develop the structure in early larval development via invaginations of the flattened planula structure. Further observations of the siphonophore species Nanomia bijuga indicate that the pneumatophore feature potentially also functions to sense pressure changes and regulate chemotaxis in some species.
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 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. Colonial movement is determined by individual nectophores of all developmental stages. The smaller individuals are concentrated towards the top of the siphonophore, and their function is turning and adjusting the orientation of the colony. Individuals will get larger the older they are. The larger individuals are located at the base of the colony, and their main function is thrust propulsion. These larger individuals are important in attaining the maximum speed of the colony. Every individual is key to the movement of the aggregate colony, and understanding their organization may allow us to make advances in our own multi-jet propulsion vehicles. The colonial organization of siphonophores, particularly in Nanomia bijuga confers evolutionary advantages. A large number of concentrated individuals allows for redundancy. This means that even if some individual nectophores become functionally compromised, their role is bypassed so the colony as a whole is not negatively affected. The velum, a thin band of tissue surrounding the opening of the jet, also plays a role in swimming patterns, shown specifically through research done on the previous mentioned species N. bijuga. The velum becomes smaller and more circular during times of forward propulsion compared to a large velum that is seen during refill periods. Additionally, the position of the velum changes with swimming behaviors; the velum is curved downward in times of jetting, but during refill, the velum is moved back into the nectophore. The siphonophore Namonia bijuga also practices diel vertical migration, as it remains in the deep-sea during the day but rises during the night.
Predation and feeding Siphonophores are predatory carnivores. 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. Species with large gastrozooids are capable of consuming a broad range of prey sizes. In one species of siphonophore, Rosacea cymbiformis, digestive experiments showed that ingested prey remained in gastrozooids for at least 8 hours before egestion. Like other Cnidaria, many siphonophore species exhibit nematocyst stinging capsules on branches of their tentacles called tentilla. The nematocysts are arranged in dense batteries on the side of the tentilla. When the siphonophore encounters potential prey, their tentillum react to where the tentacles create a net by transforming their shape around the prey. The nematocysts then shoot millions of paralyzing, and sometimes fatal, toxin molecules at the trapped prey which is then transferred to the proper location for digestion. Some species of siphonophores use aggressive mimicry by using bioluminescent light so the prey cannot properly identify the predator.
There are four types of nematocysts in siphonophore tentilla: heteronemes, haplonemes, desmonemes, and rhopalonemes. Heteronemes are the largest nematocysts and are spines on a shaft close to tubules attached to the center of the siphonophore. Haplonemes have open-tipped tubules with spines, but no distinct shaft. This is the most common nematocyst among siphonophores. Desmonemes do not have spines but instead there are adhesive properties on the tubules to hold onto prey. Rhopalonemes are nematocysts with wide tubules for prey.
Due to the scarcity of food in the deep sea environment, a majority of siphonophore species function in a sit-and-wait tactic for food.Dunn, Casey (2005). "Siphonophores". Retrieved 2008-07-08. 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. The Siphonophore Apolemia uvaria has the ability to break into small fragments to break into fish cages for food.