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Phylogeny of Medusozoa and the evolution of cnidarian life cycles
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Abstract To investigate the evolution of cnidarian life cycles, data from the small subunit of the ribosome are used to derive a phylogenetic hypothesis for Medusozoa. These data indicate that Cnidaria is monophyletic and composed of Anthozoa and Medusozoa. While Cubozoa and Hydrozoa are well supported clades, Scyphozoa appears to be paraphyletic. Stauromedusae is possibly the sister group of either Cubozoa or all other medusozoans. The phylogenetic results suggest that: the polyp probably preceded the medusa in the evolution of Cnidaria; within Hydrozoa, medusa development involving the entocodon is ancestral; within Trachylina, the polyp was lost and subsequently regained in the parasitic narcomedusans; within Siphonophorae, the float originated prior to swimming bells; stauromedusans are not likely to be descended from ancestors that produced medusae by strobilation; and cubozoan polyps are simplified from those of their ancestors, which possessed polyps with gastric septa and four mesogleal muscle bands and peristomial pits.
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“The absence of precambrian planktonic metazoans is consistent with our Early Cambrian estimate for the origin of the last common ancestor of living cnidarians, a population of animals whose life cycle lacked a medusa stage and thus was entirely benthic (49).”
From Estimating metazoan divergence times with a molecular clock (2004) · cites this paper as [49]
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“Based on morphological and 18S ribosomal RNA data, the Anthozoa appear to be the most ancestral class within the phylum (Bridge et al., 1995;Collins, 2002).”
From Evolutionary aspects of developmentally regulated helix-loop-helix transcription factors in striated muscle of jellyfish (2003) · cites this paper as Collins, 2002
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“Westfall’s (1965) hypothesis was corroborated by genetic evidence, which supports the monophyly of a clade that includes Scyphozoa, Cubozoa, and Hydrozoa (Bridge et al., 1995; Collins, 2002).”
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Schüler, A., Schmitz, G., Reft, A., Özbek, S., Thurm, U., & Bornberg-Bauer, E. (2015) · Genome Biology and Evolution · doi:10.1093/gbe/evv091 · Cited by 51
“Note that this interpretation is independent of whether sponges form a monophyletic or paraphyletic group (Collins 2002; Jackson et al. 2007; Sperling et al. 2009; Erwin et al. 2011) but places Placozoa as the outmost group within all other Metazoa, and sponges as an outgroup to this common clade…”
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“Moreover, Trichoplax has been grouped with ctenophores, as a sister group of cnidarians (Odorico and Miller 1997) and, most recently, as a sister group of the bilaterians, to the exclusion of cnidarians, ctenophores and sponges (Collins 1998, 2002).”
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“The two hydrozoan sister-groups are present in both molecular studies using 18S rRNA (Collins 2000, 2002), 16S rRNA (e.g. Collins et al. 2005), 28S rRNA data (Collins et al. 2006) or a combined dataset of the three markers (Cartwright et al. 2008) and studies based on morphology and life history…”
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“1A, 2 [Hyman 1940; Bridge et al. 1992; Collins 2002; Collins et al. 2006]).”
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“The possession of a free-living medusa is generally considered to be synapomorphic for Medusozoa (Collins, 2002; Marques & Collins, 2004) and in many instances it has been secondarily lost through paedomorphosis, with a subsequent radiation of the more benthic descendants (Boero & Sarà, 1987).”
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“…relationships among their four major groups of living medusozoans (i.e. hydrozoans, staurozoans, scyphozoans, and cubozoans) are still contentious (Collins, 2002; Collins, 2009; Collins et al., 2006; Kayal et al., 2013; Marques and Collins, 2004; Thiel, 1966; Uchida, 1973; Van Iten et al., 2006;…”
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“Until recently, the phylogenetic study of the diploblastic taxa at higher taxonomic levels was based mostly on the sequences of 28S and 18S rDNA (Medina et al. 2001; Collins 2002; Borchiellini et al. 2004).”
Pang, K., Matus, D. Q., & Martindale, M. Q. (2004) · Development Genes and Evolution · doi:10.1007/s00427-004-0383-7 · Cited by 40
“The NvCOE gene is basal relative to all other metazoan COE orthologues, consistent with the likely position of cnidarians as the sister taxa to the bilaterians (Collins 2002; Medina et al. 2001; Wainright et al. 1993).”
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“In addition, recent molecular analyses suggested that the monophyletic Cubozoa is a sister clade to the Scyphozoa (Collins 2002; Collins et al. 2006, Daly et al. 2007).”
BackgroundPhylogenetic position of Placozoa based on large subunit (LSU) and small subunit (SSU) rRNA genes
da Silva, F. B., Muschner, V. C., & Bonatto, S. L. (2007) · Genetics and Molecular Biology · doi:10.1590/s1415-47572007000100022 · Cited by 42
“A close relationship between Placozoa and Bilateria (and of both of these with Cnidaria) has also been suggested by some studies (Collins, 1998; Collins, 2002; Anderson and Swofford, 2004).”
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Cunha, A. F., Collins, A. G., & Marques, A. C. (2016) · Molecular Phylogenetics and Evolution · doi:10.1016/j.ympev.2016.09.012 · Cited by 40
“Most of the sequences assigned to Campanulariidae and available in Genbank were considered in the analysis, including those from Collins (2002), Collins et al. (2005, 2006), Govindarajan et al. (2006), Evans et al. (2008), Leclère et al. (2009), Ortman et al. (2010), Peña Cantero et al. (2010),…”
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“However, according to recent molecular and morphological phylogenies [5,6] the Placozoa branch off after the ctenophores and sometimes the cnidarians, both of which are morphologically much more complex.”
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BackgroundMolecular phylogenetic study in genus Hydra
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“The three latter, the Medusozoa, have a life cycle that includes polyp and medusa forms, whereas the members of Anthozoa only have polyp forms (Collins, 2002; Martindale et al., 2002).”
BackgroundJellyfishes—Significant Marine Resources with Potential in the Wound-Healing Process: A Review
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“The life cycle evolution of medusae in Meduzoa has been reported [90].”
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“1993; Waggoner 1996) or include enough morphological characters to enable inference of the position of extinct taxa (Jenner & Schram 1999; Giribet et al. 2001; Collins 2002; Jacobs et al. 2005; Collins et al. 2006); use of parsimony and of an explicit phylogeny might therefore yield additional data.”
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“(Left) A consensus phylogeny of the Cnidaria (based on [9,10,14], see [15] for an opposing topology).”
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Winkel, K. D., Hawdon, G. M., Fenner, P. J., Gershwin, L., Collins, A. G., & Tibballs, J. (2003) · Journal of Toxicology Toxin Reviews · doi:10.1081/txr-120019024 · Cited by 29
“The results of a recent phylogenetic analysis of cubozoan ribosomal gene sequences is summarised in Figure 2 (Collins, 2002).”
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“In phylogenetic analyses of cnidarians, it has been common for Anthozoa to be placed as a sister taxon to Medusozoa, a hypothesis supported by morphology and rDNA sequences (Collins, 2002; Daly et al., 2007).”
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“…the apical half of the metamorphosed stauromedusa (calyx) has characters similar to those of adult scyphomedusae and cubomedusae, such as hollow structures of tentacular origin (rhopalioids/rhopalia), circular coronal muscle, gastric filaments, and gonads (Collins, 2002; Collins et al., 2006).”
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“…apical half of the metamorphosed stauromedusa bears similar characters to those present in adult medusae of scyphozoans and cubozoans, e.g., hollow structures of tentacular origin (rhopalioids/rhopalia), circular coronal muscles, gastric filaments, and gonads (Collins, 2002; Collins et al., 2006).”
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Tiemann, H., & Jarms, G. (2009) · Marine Biology · doi:10.1007/s00227-009-1338-3 · Cited by 24
Internal Microanatomy and Zoological Affinity of the Early CambrianOlivooides
Han, J., Li, G., Shin, K., Ou, Q., Toshino, S., Wang, X., Xiaoguang, Y., Kentaro, U., Hoshino, M., Sasaki, O., et al. (2016) · Acta Geologica Sinica - English Edition · doi:10.1111/1755-6724.12641 · Cited by 24
Origin and Evolution of the First Nervous System
Lichtneckert, R., & Reichert, H. (2007) · Elsevier eBooks · doi:10.1016/b0-12-370878-8/00157-9 · Cited by 23
Taxonomy and phylogeny of Australian cubozoa
Gershwin, L. (2026) · ResearchOnline at James Cook University (James Cook University) · doi:10.25903/6b6r-bq97 · Cited by 25
Riccio, G., Nuzzo, G., Zazo, G., Coppola, D., Senese, G., Romano, L., Costantini, M., Ruocco, N., Bertolino, M., Fontana, A., et al. (2021) · Marine Drugs · doi:10.3390/md19080459 · Cited by 23
“for 18S and 28S, a denaturation step at 95 ◦C for 2 min, [35 cycles denaturation step at 95 ◦C for 1 min, annealing step at 52 ◦C (18S1/18S2; [48]), 55 ◦C (18S-AF/18S-BR, NL4F/NL4R; [49,50]), 57 ◦C (C2/D2; [51]), for 1 min and 72 ◦C of primer extension for 2 min], a final extension step at 72 ◦C for 10 min; ii.”
BackgroundSexual dimorphism in Tripedaliidae (Conant 1897) (Cnidaria, Cubozoa, Carybdeida)
Straehler-Pohl, I., Garm, A. L., & Morandini, A. C. (2014) · Zootaxa · doi:10.11646/zootaxa.3785.4.3 · Cited by 23
Pontin, D. R., & Cruickshank, R. H. (2012) · Hydrobiologia · doi:10.1007/s10750-011-0994-8 · Cited by 21
“For example, Collins (2002), Mandojana (1990), Yanagihara et al. (2002) and Alam et al. (2002) all use the name P. utriculus in their papers.”
BackgroundMethodEarth's oldest jellyfish strandings: a unique taphonomic window or just another day at the beach?
Sappenfield, A., Tarhan, L. G., & Droser, M. L. (2016) · Geological Magazine · doi:10.1017/s0016756816000443 · Cited by 24
“…are considered among the oldest and most widespread macrofaunal zooplankton groups in modern oceans (Collins, 2002; Peterson & Butterfield, 2005; Collins et al. 2006) and yet, despite the abundance, ubiquity and deep evolutionary history of medusozoans, the fossil record of this subphylum is…”
BackgroundIntegrated Evolution of Cnidarians and Oceanic Geochemistry Before and During the Cambrian Explosion
Han, J., Zhang, X., & Komiya, T. (2016) · Springer eBooks · doi:10.1007/978-3-319-31305-4_2 · Cited by 23
Fahey, B., Larroux, C., Woodcroft, B. J., & Degnan, B. M. (2008) · Biological Bulletin · doi:10.2307/25470664 · Cited by 23
Global Genomics of Man-O'-War (Physalia) Reveals Ocean Surface Biodiversity
Church, S. H., Abedon, R. B., Ahuja, N., Anthony, C. J., Destanović, D., Ramirez, D. A., Rojas, L. M., Albinsson, M. E., Trasobares, I. Á., Bergemann, R., et al. (2024) · Current Biology · doi:10.1016/j.cub.2025.05.066 · Cited by 23
Alzugaray, M. E., Hernández‐Martínez, S., & Ronderos, J. R. (2016) · Peptides · doi:10.1016/j.peptides.2016.05.011 · Cited by 22
“vulgaris (Hydrozoa), and in fact, it has been proposed that Anthozoa is the group of cnidarians more closely related to bilaterians [10, 22].”
Schuchert, P., & Reiswig, H. M. (2006) · Canadian Journal of Zoology · doi:10.1139/z06-031 · Cited by 22
“Although the latter could in fact be only a subgroup of the Anthoathecata (Schuchert 1996; Collins 2002), their extraordinary complex organization makes them an unlikely candidate for the closest relatives of B.”
BackgroundAn Overview of the Medusozoa from the Southwestern Atlantic
Schiariti, A., Dutto, M. S., Morandini, A. C., Nagata, R. M., Pereyra, D. Y., Puente-Tapia, F. A., Briz, L. M. D., & Genzano, G. (2018) · Springer eBooks · doi:10.1007/978-3-319-77869-3_19 · Cited by 22
Pruski, S., & Miglietta, M. P. (2019) · PeerJ · doi:10.7717/peerj.7848 · Cited by 21
Hydrozoans, scyphozoans, larvaceans and ctenophores observed in situ at hadal depths
Jamieson, A. J., & Linley, T. D. (2020) · Journal of Plankton Research · doi:10.1093/plankt/fbaa062 · Cited by 21
“These taxonomic groups are often associated with deep pelagic environments with diverse and often cryptic speciation spanning the bathyal (200–3000 m) and abyssal (3000–6000 m) zones (Haddock and Case, 1999; Collins, 2002; Hopcroft, 2005; Matsumoto et al., 2020).”
BackgroundThe state of Medusozoa genomics: current evidence and future challenges
Santander, M. D., Maronna, M. M., Ryan, J. F., & Andrade, S. C. S. (2022) · GigaScience · doi:10.1093/gigascience/giac036 · Cited by 21
“Most medusozoan life cycles are characterized by the succession of different stages, including a larval, benthic asexually reproducing polyp stage and a sexually reproducing jellyfish stage [6, 7].”
BackgroundLight modulated cnidocyte discharge predates the origins of eyes in Cnidaria
Picciani, N., Kerlin, J. R., Jindrich, K., Hensley, N. M., Gold, D. A., & Oakley, T. H. (2021) · Ecology and Evolution · doi:10.1002/ece3.7280 · Cited by 20
A Cnidarian Phylogenomic Tree Fitted With Hundreds of 18S Leaves
DeBiasse, M. B., Buckenmeyer, A., Macrander, J., Babonis, L. S., Bentlage, B., Cartwright, P., Prada, C., Reitzel, A. M., Stampar, S. N., Collins, A. G., et al. (2024) · Bulletin of the Society of Systematic Biologists · doi:10.18061/bssb.v3i2.9267 · Cited by 20
Self-citationSchwarz, R. S., Bosch, T. C. G., & Cadavid, L. F. (2008) · Developmental & Comparative Immunology · doi:10.1016/j.dci.2008.03.007 · Cited by 19
“Within the Cnidaria, class Anthozoa and Hydrozoa are the most ancestral of the four currently recognized classes [4,5].”
BackgroundCyathophycus and the origin of demosponges
Botting, J. P. (2003) · Lethaia · doi:10.1080/00241160310006385 · Cited by 19
Miranda, T. P., Genzano, G., & Marques, A. C. (2015) · Zootaxa · doi:10.11646/zootaxa.4033.4.2 · Cited by 19
“Since then, many phylogenetic classifications for the more inclusive taxa of Hydrozoa were proposed (Collins 2000, 2002; Marques 2001; Marques & Collins 2004; Collins et al. 2005, 2006, 2008; Cartwright et al. 2008; Leclère et al. 2009; Cartwright & Nawrocki 2010; Nawrocki et al. 2010).”
BackgroundExpression of aGsx parahox gene,Cnox-2, in colony ontogeny inHydractinia (Cnidaria: Hydrozoa)
Cartwright, P., Schierwater, B., & Buss, L. W. (2006) · Journal of Experimental Zoology Part B Molecular and Developmental Evolution · doi:10.1002/jez.b.21106 · Cited by 19
Nme family of proteins—clues from simple animals
Ćetković, H., Perina, D., Harcet, M., Mikoč, A., & Bosnar, M. H. (2014) · Naunyn-Schmiedeberg s Archives of Pharmacology · doi:10.1007/s00210-014-1017-x · Cited by 18
“(Collins 2002).”
Yamamori, L., Okuizumi, K., Sato, C., Ikeda, S., & Toyohara, H. (2017) · ZOOLOGICAL SCIENCE · doi:10.2108/zs160161 · Cited by 18
Martindale, M. Q., & Henry, J. Q. (2015) · Evolutionary Developmental Biology of Invertebrates 1 · doi:10.1007/978-3-7091-1862-7_8 · Cited by 18
Molecular Characterisation, Evolution and Expression of Hypoxia-Inducible Factor in Aurelia sp.1
Wang, G., Yu, Z., Zhen, Y., Mi, T., Shi, Y., Wang, J., Wang, M., & Sun, S. (2014) · PLoS ONE · doi:10.1371/journal.pone.0100057 · Cited by 18
“The above description indicates that the polyps most likely preceded the medusa in Cnidarian evolution [20].”
BackgroundHypša, V. (2006) · International Journal of Parasitology · doi:10.1016/j.ijpara.2005.10.010 · Cited by 19
Nicosia, A., Maggio, T., Mazzola, S., Gianguzza, F., Cuttitta, A., & Costa, S. (2014) · PLoS ONE · doi:10.1371/journal.pone.0105908 · Cited by 17
“The Medusozoa comprises three classes Scyphozoa (true jellyfishes), Cubozoa (box jellyfishes) and Hydrozoa (hydras and hydromedusae).”
BackgroundCell Proliferation in Cubozoan Jellyfish Tripedalia cystophora and Alatina moseri
Gurská, D., & Garm, A. L. (2014) · PLoS ONE · doi:10.1371/journal.pone.0102628 · Cited by 19
“Among cnidarians only cubozoans undergo a complete metamorphosis from polyp to medusa in that the entire polyp turns into a single medusa [1,2].”
BackgroundEvolution of Metazoan Life Cycles and the Origin of Pelagic Larvae
Malakhov, V. V., Bogomolova, E., Kuzmina, T. V., & Temereva, E. N. (2019) · Russian Journal of Developmental Biology · doi:10.1134/s1062360419060043 · Cited by 17
“It should be noted that, according to modern ideas, Medusozoa (including Hydrozoa) is an apomorphic group of Cnidaria, in the life cycle of which an additional medusa stage appeared (Collins, 2002; Marques and Collins, 2004; Collins et al., 2006).”
BackgroundInfluentialAbouna, S., Gonzalez‐Rizzo, S., Grimonprez, A., & Gros, O. (2015) · PLoS ONE · doi:10.1371/journal.pone.0127625 · Cited by 17
Jacobs, D. K. (2003) · Integrative and Comparative Biology · doi:10.1093/icb/43.1.11 · Cited by 17
“Small ribosomal subunit (18S) phylogenics place Anthozoa the class of Cnidaria lacking medusae, as the basal branch of living cnidarians (e.g., Collins, 2002).”
Back to the roots, Part 1—early life cycle data of Rhopaliophora (Scyphozoa, Cubozoa and Staurozoa)
Straehler-Pohl, I., & Jarms, G. (2022) · Plankton and Benthos Research · doi:10.3800/pbr.17.1 · Cited by 16
Lieberman, B. S., Kurkewicz, R., Shinogle, H. E., Kimmig, J., & MacGabhann, B. A. (2017) · PeerJ · doi:10.7717/peerj.3312 · Cited by 16
Microbial diversity in Mediterranean sponges as revealed by metataxonomic analysis
Ruocco, N., Esposito, R., Zagami, G., Bertolino, M., Matteo, S. D., Sonnessa, M., Andreani, F., Crispi, S., Zupo, V., & Costantini, M. (2021) · Scientific Reports · doi:10.1038/s41598-021-00713-9 · Cited by 16
Metadata
- Authors
- Collins, A. G.
- Affiliations
- University of California, Berkeley (US)
- Corresponding author
- Collins, A. G.
- ORCID
- Collins, A. G.
- Venue
- Journal of Evolutionary Biology
- Publisher
- Oxford University Press (OUP)
- Published
- 2002-05-01
- Volume
- 15
- Issue
- 3
- Pages
- 418-432
- ISSN
- 1010-061X (ISSN-L)
- 1420-9101
- OpenAlex
- W2098787235
- Cited by
- 355
- References
- 62
- Citations by year
- 2026: 11 · 2025: 8 · 2024: 9 · 2023: 12 · 2022: 16 · 2021: 10 · 2020: 6 · 2019: 13 · 2018: 15 · 2017: 19 · 2016: 21 · 2015: 27 · 2014: 15 · 2013: 9 · 2012: 7
- FWCI
- 17.71
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- Journal of Evolutionary Biology (publishedVersion)
- Funders
- National Science Foundation (9814845)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Animal Science and Zoology › Marine Invertebrate Physiology and Ecology
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Protist diversity and phylogeny
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Invertebrate Taxonomy and Ecology
- Keywords
- Medusozoa, Anthozoa, Cubozoa, Hydrozoa, Scyphozoa, Siphonophorae, strobilation, monophyly
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