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Annotated Bibliography
Haddock, Steven H. D., Dunn, Casey W., Pugh, Philip R., Schnitzler, Christine E. (2005). Bioluminescent and Red-Fluorescent Lures in a Deep-Sea Siphonophore. Science, vol. 309, p. 263. DOI: 10.1126/science.1110441
 * Most siphonophores are bioluminescent, and they often use this chemically-induced light as a defensive strategy against predators. Siphonophores are also effective predators themselves, yet they are extremely fragile.
 * Data was collected from three siphonophores belonging to the genus, Erenna, at depths between 1,600 - 2,300 meters. It revealed that Erenna feeds on fish, and researchers hypothesize that this non-visual species does so by emitting red-fluorescence in their tentilla along with a flicking pattern to lure fish.
 * There is a view that deep-sea organisms cannot detect long wavelengths (red light has a wavelength of 680 nm). If this is the case, then it is not possible for the fish to be lured by Erenna and there must be another explanation. However, the deep-sea remains largely unexplored and we should not discard red-light sensitivity.

Purcell, Jennifer E. (1980). Influence of Siphonophore Behavior upon Their Natural Diets: Evidence for Aggressive Mimicry. Science, vol. 209, pp. 1045-1047. DOI: 10.1126/science.209.4460.1045


 * The diets of siphonophore species within 11 genera were studied at depths between 0 - 25 m. In general, the diets of strong swimming species consist of small prey while the diets of weak swimming species consist of larger prey.
 * Strong swimmers change locations often in order to encounter high-density prey.
 * Weak swimmers are relatively inactive, lowering their chances of encountering prey while simultaneously saving energy.
 * Species with large gastrozooids consume a broad range of prey sizes.
 * To capture prey, siphonophores use their tentacles as “webs” that prey swim into. Within the tentacles, there are millions of toxic nematocysts (specialized cells in the tentilla) that sting and paralyze their prey.
 * Two siphonophore species attract relatively large prey by mimicking small zooplankton. These species possess nematocyst batteries that help deceive and poison the prey.

Palma, Sergio, Cabello, Fabiola, Silva, Nelson, Canepa, Antonio (2017). Siphonophores of the Chiloé Inland Sea: biodiversity, spatial distribution, and environmental association. Marine Biodiversity vol. 48, pp. 1731-1742. DOI: 10.1007/s12526-017-0662-y.


 * For this study, nine siphonophore species were identified in the Chiloé Inland Sea. The purpose was to study their abundance, distribution, and ecological impact in the area.
 * In general, siphonophore densities are most concentrated in the north micro basin, which contains estuarine waters (which according to the text consist of “greater vertical stratification, lower temperatures and salinity, and a higher concentration of dissolved oxygen”) at depths of 0 - 50 m. In the south micro basin, they are mostly found at depths > 50 m.
 * Siphonophores are predators with important ecological roles. According to the text, they consume a wide range of prey including zooplankton, which may be of commercial interest. As an example, Muggiaea atlantica has negative effects on salmon farming.
 * In Chile, siphonophores have been studied extensively in regards to their taxonomy and the distribution of the most abundant and the most ecologically important species. 55 out of 175 siphonophore species have been observed along the Chilean coast, and 15 have been observed in the Chiloé Inland Sea.

Araya, Juan Francisco, Aliaga, Juan Antonio, Esther Araya, Marta (2016). On the distribution of Physalia physalis (Hydrozoa: Physaliidae) in Chile. Marine Biodiversity vol. 46 pp. 731-735. DOI: 10.1007/s12526-015-0417-6


 * Several colonies of Physalia physalis, otherwise known as the Portuguese Man o’ War, washed ashore in the Atacama Region of Chile in 2015. This abundance likely occurred due to El Niño Southern Oscillation events, and it incentivized a study of the species based on new findings.
 * This species is mostly found in tropical and subtropical regions, where the ocean is warm.
 * Physalia physalis is almost transparent, it floats in the surface of the ocean because of its air bladder, and it is able to navigate because of its erectile sailing crest. This species has thin tentacles, dactylozooids, that according to the text are “capable of discharging thousands of cnidae, which depend on mechanical and chemical stimuli, producing acute envenoming in humans and even death caused by vasomotor dysfunction and collapse”.

Costello, John H., Colin, Sean P., Gemmell, Brad J., Dabiri, John O., Sutherland, Kelly R. (2015). Multi-jet propulsion organized by clonal development in a colonial siphonophore. Nature Communications, vol. 6, 8185. DOI: 10.1038/ncomms9158


 * Siphonophores are unique in the sense that they are made up of clonal individuals, or nectophores, that form by budding and are genetically equal. Nectophores join in a complex aggregate colony organization.
 * For this study, the nectophores of the species Nanomia bijuga were observed. In general, young and weak nectophores “dominate torque production for turning” while older and more powerful nectophores “contribute predominantly to forward thrust production”. 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 nectophores of Nanomia bijuga allow for the species to swim forwards and reverse, to spiral and contract, and to swim in many other complex ways. Nanomia bijuga practices vertical migration every day, as it remains in the deep during the day but rises during the night.
 * According to this text, the aggregate colony organization of siphonophores is an evolutionary advantage. It has allowed for the organism to grow extensively. Additionally, if and when individual nectophores suffer damage, their role is bypassed by other nectophores (even when every nectophore - weak or powerful - is essential to the mechanism of swimming).

Mapstone, Gillian M. (2014). Global Diversity and Review of Siphonophorae (Cnidaria: Hydrozoa). Public Library of Science, vol. 9, 2. DOI: 10.1371/journal.pone.0087737


 * This 37-page paper is a summary of the history of siphonophores. The author aims to cover the first discovery of siphonophores in 1785 (Physalia physalis) until the most recent findings, which includes their overall distribution, abundance, and body-plans (e.g. their tentilla can incorporate toxic nematocysts and/or serve as lures that emit red-fluorescence).
 * There is no fossil record of siphonophores, even when they have evolved and developed for an extensive period (its phylum, Cnidaria, dates back to 640 million years).
 * Smaller species live in the epipelagic zone where they mostly feed on zooplankton, while larger species live in the mesopelagic zone where they mostly feed on larger prey. They hunt by casting “webs” with their tentacles that essentially traps and poisons prey.
 * 91% of siphonophores are bioluminescent, though most use this for defensive purposes. However, the tentilla of some larger deep-sea siphonophores emit bioluminescence. This light attracts prey as it may resemble smaller organisms such as zooplankton and copepods. For example, the species Erenna resides between 1600 - 2300 m, and it has adapted in a way that allows it to produce red-fluorescence in a rhythmic pattern that attracts deep-sea fish.
 * This paper includes a variety of pictures, diagrams, charts and phylogenetic trees that are useful to further understand the nature of siphonophores

Ideas for Editing Siphonophorae
Bioluminescence Section


 * Original Text:
 * 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 in the genus Erenna are thought to use their bioluminescent capability as a lure to attract fish. This genus is one of the few to prey on fish rather than crustaceans. The bioluminescent organs on these individuals flicker, and thus it has been concluded that they use bioluminescence to attract prey. In addition to their use of bioluminescence, organisms in the genus Erenna use red fluorescence to attract prey.
 * Change Original Text:
 * "Siphonophores of the deep-sea genus Erenna (found at depths between 1,600-2,300 meters) are thought to use their bioluminescence capability for offense too, as a lure to attract fish."
 * "The bioluminescent organs (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 bioluminescence as a lure to attract prey.
 * Add Text:
 * "There is a view that deep-sea organisms can not detect long wavelengths, and red light has a 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."
 * Cite: Haddock, Steven H. D., Dunn, Casey W., Pugh, Philip R., Schnitzler, Christine E. (2005). Bioluminescent and Red-Fluorescent Lures in a Deep-Sea Siphonophore. Science, vol. 309, p. 263. DOI: 10.1126/science.1110441
 * Cite: Mapstone, Gillian M. (2014). Global Diversity and Review of Siphonophorae (Cnidaria: Hydrozoa). Public Library of Science, vol. 9, 2. DOI: 10.1371/journal.pone.0087737

Predation and feeding Section


 * Original Text:
 * Siphonophores are predatory carnivores. Their diets consist of a variety of copepods, small crustaceans, and small fish. 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. Similar to many other organisms in the phylum of 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, they utilize their 30–50 cm (12–20 in) tentacles to create a net around the prey. The nematocysts then shoot paralyzing, and sometimes fatal, toxins at the trapped prey which is then transferred to the proper location for digestion.
 * Change Original Text:
 * "Their diets consist of a variety of copepods, small crustaceans, and small fish. Generally, the diets of strong swimming siphonophores consist of smaller prey, and the diets of weak swimming siphonophores consist of larger prey. 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."
 * "The nematocysts then shoot millions of paralyzing, and sometimes fatal, toxins molecules at the trapped prey which is then transferred to the proper location for digestion." (would I have to use a citation for "millions"?)


 * Cite: Purcell, Jennifer E. (1980). Influence of Siphonophore Behavior upon Their Natural Diets: Evidence for Aggressive Mimicry. Science, vol. 209, pp. 1045-1047. DOI: 10.1126/science.209.4460.1045

Movement Section


 * Original Text:
 * Siphonophores use a method of locomotion similar to jet propulsion. A nectophore is a gathering of many siphonophores, and depending on where each individual siphonophore is positioned within the nectophore, their function differs. Colonial movement is determined by individual siphonophores of all developmental stages. The smaller individuals are concentrated towards the top of the nectophore, 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. The colonial organization of siphonophores, particularly in Nanomia bijuga confers evolutionary advantages. A large amount of concentrated individuals allows for redundancy. This means that even if some individual siphonophores become functionally compromised, the colony as a whole is not negatively affected.
 * There's a major mistake in this paragraph, located in the second sentence: "A nectophore is a gathering of many siphonophores, and depending on where each individual siphonophore is positioned within the nectophore, their function differs." Instead, it should say "A siphonophore is a gathering of many nectophores, and depending on where each individual nectophore is positioned within the nectophore, their function differs."


 * Change Original Text:
 * "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 amount 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."
 * Nanomia bijuga also practices diel vertical migration, as it remains in the deep during the day but rises during the night. This could potentially contribute to the Movement section.
 * Cite: Costello, John H., Colin, Sean P., Gemmell, Brad J., Dabiri, John O., Sutherland, Kelly R. (2015). Multi-jet propulsion organized by clonal development in a colonial siphonophore. Nature Communications, vol. 6, 8185. DOI: 10.1038/ncomms9158

Discovery Section


 * Original Text:
 * Carl Linnaeus discovered and described the first siphonophore, the Portuguese man o' war, in 1758. The discovery rate of siphonophore species was slow in the 18th century, as only four additional species were found. During the 19th century, 56 new species were observed due to research voyages conducted by European powers. The majority of new species found during this time period were collected in coastal, surface waters. During the HMS Challenger expedition, various species of siphonophores were collected. Ernst Haeckel attempted to conduct a write up of all of the species of siphonophores collected on this expedition. He introduced 46 "new species"; however, his work was heavily critiqued because some of the species that he identified were eventually found not to be siphonophores. Nonetheless, some of his descriptions and figures (pictured below) are considered useful by modern biologists. A rate of about 10 new species discoveries per decade was observed during the 20th century. Considered the most important researcher of siphonophores, A. K. Totton introduced 23 new species of siphonophores during the mid 20th century. On April 6, 2020 the Schmidt Ocean Institute announced the discovery of a giant Apolemia siphonophore in submarine canyons near Ningaloo Coast, measuring 15 m (49 ft) diameter with a ring approximately 47 m (154 ft) long, possibly the largest siphonophore ever recorded.
 * Change Original Text:
 * "There is no fossil record of siphonophores, though they have evolved and adapted for an extensive time period. Their phylum, Cnidaria, is an ancient lineage that dates back to c. 640 million years ago."
 * Cite: Mapstone, Gillian M. (2014). Global Diversity and Review of Siphonophorae (Cnidaria: Hydrozoa). Public Library of Science, vol. 9, 2. DOI: 10.1371/journal.pone.0087737

Distribution and habitat Section

Ideas for Future Improvements
 * Original Text:
 * 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. 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 and/or an area of the ocean.
 * Change Original Text:
 * "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."
 * Cite: Mapstone, Gillian M. (2014). Global Diversity and Review of Siphonophorae (Cnidaria: Hydrozoa). Public Library of Science, vol. 9, 2. DOI: 10.1371/journal.pone.0087737


 * Add a section discussing siphonophore tentilla and nematocysts?
 * Add a section about the presence and environmental impact of siphonophores on the Chilean coast?
 * Add a section about siphonophore ecological r oles (e.g. Muggi aea atlantica's negative effect on salmon farming)?
 * Add more on the siphonophore Portuguese Man o ' War (Physalia physalis) (found in tropical/subtropical regions, floats on the surface due to air bladder, navigates due to erectile sailing crest, dactylzooids are capable of discharging venomous toxins that can potentially cause vasomotor dysfunction and collapse, etc.)?
 * Add more photos to make the text more engaging?
 * Add the date that siphonophores first evolved?