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A colony of the rare hydrozoan siphonophore Lilyopsis Chun, 1885, was observed for the first time in shallow water in False Bay, South Africa, swimming amongst kelp. A study of a high-quality image of this individual found it to share some characters with the prayine prayid L. fluoracantha Haddock, Dunn & Pugh, 2005, so far known only from Monterey Bay, California, in the eastern Pacific. No Lilyopsis species has previously been reliably identified from either the South Atlantic or the Indian Ocean, so this record represents an expansion of the known worldwide distribution for this genus.

This report documents the first record of the genera Eucheilota and Mitrocomella and species Porpita porpita and Physalia physalis in Bali, Indonesia, based on observed occurrences in different times and sites. The coincidence of the annual stranding of Physalia physalis in the east Bali and south Java coasts during the monsoon periods in Indonesia suggests a link with the upwelling events in the areas. However, more work is needed to analyse this phenomenon and study the occurrences of other Hydromedusae due to the limited data on hydrozoans in Indonesian waters. Overall, this report provides primary data to contribute to the general understanding of the biodiversity of marine organisms in Indonesia.

Orientador: Prof. Dr. Otto Müller Patrão de Oliveira Tese (doutorado) - Universidade Federal do ABC, Programa de Pós-Graduação em Evolução e Diversidade, Santo André, 2021. Os sifonóforos são um grupo de hidrozoários (Cnidaria, Medusozoa) de grande importância no ambiente pelágico marinho, sendo potencialmente um dos principais predadores do zooplâncton presente nesta área. Apesar disso, estudos sobre esses organismos são escassos na literatura. Este trabalho apresenta descrições morfológicas para todas as espécies válidas de sifonóforos. O material examinado é proveniente de diversas instituições ao redor do mundo: Laboratório de Zooplâncton Gelatinoso da Universidade Federal do ABC (Brasil); Smithsonian National Museum of Natural History (Estados Unidos); Yale Peabody Museum of Natural History (Estados Unidos); Natural History Museum London (Inglaterra); University Museum of Bergen (Noruega); National Museum of Nature and Science (Japão); e Melbourne Museum (Austrália). Ao todo, 2244 espécimes, relativos a 171 de 190 espécies válidas de sifonóforos, foram fotografados, descritos, esquematizados e medidos. As demais espécies que não foram examinadas no estudo também foram descritas, com base em dados da literatura. As descrições das espécies foram feitas com base em estudos filogenéticos recentes, de forma a refletir a taxonomia atual do grupo. Adicionalmente, foram elaborados mapas de distribuição para todas as espécies válidas de sifonóforos. The siphonophores are a group of hydrozoans (Cnidaria, Medusozoa) of great importance in the marine pelagic environment, being potentially one of the main zooplankton predators present in this area. Despite that, studies about these organisms are scarce in the literature. This work presents morphological descriptions for all valid siphonophore species. The material examined comes from several institutions around the world: Laboratory of Gelatinous Zooplankton of Federal University of ABC (Brazil); Smithsonian National Museum of Natural History (United States); Yale Peabody Museum of Natural History (United States); Natural History Museum London (England); University Museum of Bergen (Norway); National Museum of Nature and Science (Japan); and Melbourne Museum (Australia). A total of 2244 specimens, relative to 171 of 190 valid siphonophore species, were photographed, described, schematized and measured. The remaining species that were not examined in the present study were also described, based on literature data. The species descriptions were made based on recent phylogenetic studies, in order to reflect the group's current taxonomy. Additionally, distribution maps of all valid siphonophore species were elaborated.

This chapter includes a review of the recent results regarding zooplankton in offshore Montenegrin waters (since 2009) and a comparison of these results with earlier surveys. The highest abundance of total zooplankton was determined at station E, close to the river Bojana, in the summer period reaching 74, 972 ind m 3. A lower density of total zooplankton was observed during the winter. The abundance of total zooplankton increased toward the southern part of Montenegrin waters and from offshore to coastal stations. Contrary to the abundance, biodiversity grew from the coastal to the offshore sites as well as from the southern to the northern part of Montenegrin waters. Thirteen groups and 126 mesozooplankton taxa were determined. Analysis of the 12-month monitoring revealed that copepods were the most dominant group with an average contribution of 79% and a maximum share of 99%. Copepods were the most diverse group with 44 determined taxa. The most dominant taxa were Onceaidae-like species with 17% share, followed by: Oithona nana (14%), Acartia (Acartiura) clausi and Euterpina acutifrons (10%), and Paracalanus parvus parvus (8.5%). Cladoceran species dominated during the summer period, but the average contribution was 10%. Noctiluca scintillans reached a maximum value of 6, 417 ind m 3 in August 2009 at A3 station. Seventeen taxa of hydromedusae and 11 species of siphonophorae were collected during the survey period. Chelophyes appendiculata was noted for the first time in Montenegrin waters. High abundances of meroplankton larvae were recorded in this area throughout the investigation period, representing 2.7% of the total mesozooplankton. Bivalve larvae generally constituted the majority of this population. Biodiversity indexes showed statistically significant difference by seasons and decreasing values from transect A to transect E but no statistical difference was found. Comparison of medians of coastal with offshore stations showed statistically significant difference for the Margalef’s index. The data presented here nevertheless suggest where limited sampling resources should be used to describe more confidently the functional role of the mesozooplankton community in the Montenegrin Adriatic offshore waters.

Hydrozoa, Scyphozoa, Larvacea (Appendicularia) and Ctenophora are not typically associated with hadal communities. Here, we report observations of these groups based on 136 benthic camera lander deployments that spanned all five oceans, encompassing 14 deep sites, culminating in >1000 h of video in the near-bottom waters between 5000 and 10 925 m. Of the Hydrozoa, trachymedusae had a maximum depth of 9066 m in the Mariana Trench, narcomedusae were recorded to a maximum depth of 7220 m in the San Cristobal Trench and a single siphonophore was seen at 7888 m in the Mariana Trench. Scyphozoans were seen as deep as 6898 m in the New Hebrides Trench. The deepest ctenophore was seen at 6037 m in the Kermadec Trench. Larvaceans were seen in the Agulhas Fracture Zone and the Puerto Rico, Kermadec, South Shetland and Java trenches, with the deepest being 7176 m in the Java Trench. None of these groups were seen in the deep Arctic or Antarctic deeper than 6000 m. Narcomedusae, siphonophorae, Scyphozoa and Ctenophora appear very rare at hadal depths, while the larvaceans and trachymedusae appear to be relatively conspicuous in the benthopelagic at hadal depths.

Rhodalia miranda Haeckel, 1888 Material examined: PD BB 2017: St. 30; St. 31. Distribution: Atlantic Ocean, Uruguay-Argentina, from 35º to 56ºS on slope (Riemann-Zurneck 1991). Remarks: Epibenthic colonial siphonophorae with demersal habit were found., Published as part of Schejter, L., Genzano, G., Pérez, C. D., Acuña, F., Cordeiro, R. T. S., Silva, R. A., Garese, A. & Bremec, C. S., 2020, Checklist of Benthic Cnidaria in the SW Atlantic Ocean (54 ºS- 56 ºS), pp. 201-239 in Zootaxa 4878 (2) on page 215, DOI: 10.11646/zootaxa.4878.2.1, http://zenodo.org/record/4424884, {"references":["Riemann-Zurneck, K. (1991) The benthic deep-water siphonophore Rhodalia miranda and other coelenterates in the south-west Atlantic: ecological and oceanographical implications. Hydrobiologia, 216 (217), 481 - 487. https: // doi. org / 10.1007 / BF 00026502"]}

In this study we provide an updated checklist of benthic Cnidaria from SW Atlantic Ocean that comprised the Marine Protected Areas Namuncurá I and II, located at Burdwood bank, and other neighbouring locations. A total of 88 taxa was recorded: 36 hydrozoans and 52 anthozoans from which 32 were octocorals, 10 scleractinian corals, 8 sea anemones and 2 zoanthids. Burdwood bank presented the highest richness considering that 87% of the recorded species inhabit this plateau or its slope. Besides some common species widely distributed in the studied sub-areas, at least 24 species represent new distributional records while few were exclusively recorded at Burdwood bank. The inventory here provided will help to identify key habitat-forming species in a complex habitat where marine animal forests and vulnerable marine ecosystems were previously detected. It will be also a very valuable tool for the management and monitoring of the sub-areas under protection.

Mikrolepidopteren gehörten seit der ersten Hälfte des 19. Jahrhunderts zum Gegenstand entomologischer Forschung in Eberswalde. Zentren der Forschung waren zunächst die Forstlichen Lehranstalten und Institute, später auch das Deutsche Entomologische Institut. Außerdem war die Eberswalder Gegend stets Anziehungspunkt für viele Mikrolepidopteren-Spezialisten, die nicht an einschlägigen wissenschaftlichen Instituten arbeiteten. In der vorgelegten Arbeit werden die Forschungsergebnisse aus der Vergangenheit regionalfaunistisch ausgewertet und durch Ergebnisse aktueller faunistischer Forschungen zu einem Verzeichnis der Microlepidoptera der Diluviallandschaften um Eberswalde ergänzt. Das Verzeichnis umfasst mit 863 Arten aus 47 Familien 57 % der im Land Brandenburg nachgewiesenen Mikrolepidopteren-Arten, darunter ein Neufund für Deutschland: Clepsis coriacanus; 13 Neufunde für das Land Brandenburg: Coleophora adjectella, Colephora sylvaticella, Coleophora ramosella, Mompha terminella, Monochroa arundinetella, Bryotropha umbrosella, Anarsia innoxiella, Telephila schmidtiellus, Cydia gemmiferana, Acrobasis legatea, Pammene herrichiana, Pammene ignorata, Schreckensteinia festaliella; und 45 Wiederfunde im Land Brandenburg. Die Autoren konnten 332 Arten erstmals im Eberswalder Raum nachweisen. Für 531 Arten wurden ältere/historische Nachweise ausgewertet. Das Vorkommen von 408 dieser Arten konnte bestätigt werden. Die Funddaten sind tabellarisch dargestellt und sowohl Naturräumen des Gebietes als auch Messtischblättern zugeordnet., Beiträge zur Entomologie = Contributions to Entomology, Bd. 70 Nr. 1 (2020)

This study compares the structure of the zooplankton community in the Bay of Bengal (BoB) and South China Sea (SCS) during the period of spring inter-monsoon, 2010. A total of 215 species of zooplankton were identified, of which 187 species were present in the BoB and 119 in the SCS. Of the taxonomic groups recorded, Copepoda was the most diverse group in all samples followed by pelagic Tunicata, Siphonophorae and Chaetognatha. Flaccisagitta enflata, Cosmocalanus darwinii, Euchaeta larva, Macrura larva and Candacia truncata were predominant both in the BoB and SCS. Moreover, the distribution of some dominant species differed regionally, such as Cypridina dentata, Pleuromamma robusta and Mesosagitta decipiens only in the BoB, and Pleuromamma gracilis, Neocalanus gracilis and Eudoxoides spiralis in the SCS. The average zooplankton abundance was 33.37 ± 7.19 ind. m−3 in the BoB and 35.08 ± 2.07 ind. m−3 in the SCS. Copepoda was one of the most abundant groups in the BoB and SCS. Based on multivariate analysis, it was possible to distinguish the zooplankton in the BoB and SCS communities at the similarity level of approximately 55%, and the dissimilarity was mainly due to C. dentata, P. robusta, M. decipiens, C. darwinii, N. gracilis and P. gracilis. The relationships between zooplankton and temperature, salinity and chlorophyll a were not statistically significant. Zooplankton community structure in the BoB and SCS was observed to be generally similar in terms of species composition and abundance, but the differences observed may be the result of species-specific geographical distribution and local hydrographic conditions.

The Tara Oceans project sampled contrasting ecosystems of the world oceans during a three-year expedition (2009-2013), collecting environmental data and plankton, from viruses to metazoans, on board the 36-metre Tara Schooner. It surveyed 210 ecosystems in 20 biogeographic provinces, collecting over 35,000 samples of seawater and plankton. Samples were later analysed using modern sequencing and state-of-the-art imaging technologies., We are keen to thank the commitment of the following people and sponsors who made this singular expedition possible: CNRS, EMBL, Genoscope/CEA, VIB, Stazione Zoologica Anton Dohrn, UNIMIB, ANR (projects POSEIDON, BIOMARKS, PROMETHEUS, and TARA-GIRUS), FWO, BIO5, Biosphere 2, agnès b., the Veolia Environment Foundation, Region Bretagne, World Courier, Illumina, Cap L'Orient, the EDF Foundation EDF Diversiterre, FRB, the Prince Albert II de Monaco Foundation, Etienne Bourgois, the Tara schooner and its captain and crew. Tara Oceans would not exist without the continuous support of the participating 23 institutes

The Tara Oceans project sampled contrasting ecosystems of the world oceans during a three-year expedition (2009-2013), collecting environmental data and plankton, from viruses to metazoans, on board the 36-metre Tara Schooner. It surveyed 210 ecosystems in 20 biogeographic provinces, collecting over 35,000 samples of seawater and plankton. Samples were later analysed using modern sequencing and state-of-the-art imaging technologies., We are keen to thank the commitment of the following people and sponsors who made this singular expedition possible: CNRS, EMBL, Genoscope/CEA, VIB, Stazione Zoologica Anton Dohrn, UNIMIB, ANR (projects POSEIDON, BIOMARKS, PROMETHEUS, and TARA-GIRUS), FWO, BIO5, Biosphere 2, agnès b., the Veolia Environment Foundation, Region Bretagne, World Courier, Illumina, Cap L'Orient, the EDF Foundation EDF Diversiterre, FRB, the Prince Albert II de Monaco Foundation, Etienne Bourgois, the Tara schooner and its captain and crew. Tara Oceans would not exist without the continuous support of the participating 23 institutes (see http://oceans.taraexpeditions.org).

Two new deep-living species of the genus Bargmannia (Siphonophorae, Physonectae, Pyrostephidae) are described based, almost exclusively, on the nectophores of single specimens collected by ROVs off the west coast of California. The nectophores of B. stenotes sp. nov. were relatively small, up to 11.4 mm in length and, in the preserved condition had very narrow nectosacs. There was a marked bend in the distal part of the preserved nectophore, resulting in the ostium opening upwards. The mature nectophores of B. profunda sp. nov. were almost twice as long as those of B. stenotes and were characterised by the fact that the lower lateral ridges did not join, distally, with the meso-lateral ones. This feature also was present in smaller nectophores of B. amoena but those of B. stenotes, at that stage, did not have a mouth-plate.

Pugh, P.R. (2019): A review of the siphonophore species mentioned in Haeckel's (1888 b) Challenger Monograph. Zootaxa 4683 (1): 1-32, DOI: https://doi.org/10.11646/zootaxa.4683.1.1

In his Challenger Monograph Haeckel (1888b) listed a total of 204 species of siphonophores and 36 species of Porpitidae (Disconectae, according to Haeckel), which are now known not to belong to the Siphonophorae. In this paper the siphonophore species have been divided into four categories: a). species that had been described previously by an acknowledged authority other than Haeckel; b). species where Haeckel changed the generic or specific name of a previously described species by another authority, and then ascribed the authority to himself; c). species that Haeckel actually described and illustrated as what he believed to be new; and d). species that Haeckel mentioned in the text as a new species, but with the description deferred to a later publication or simply not given. The validity of the forty-three species that Haeckel actually described is then discussed. A full list of all these species is given in an Appendix.

Rhizophysa filiformis (Forssk��l, 1775) Physsophora filiformis Forssk��l, 1775, p. 120; 1776, Pl. 33, fig. F; Modeer, 1789, p. 282; Gmelin, 1790, p. 3159; Lamarck, 1816, p. 478; 1840, p. 82; Eysenhardt, 1821, p. 416; Blainville, 1826, p. 151; 1834, p. 118; Bosc, 1830, p. 200; Griffiths & Pidgeon, 1834, p. 491; Rhizophysa filiformis Oken, 1815, p. 127; Lamarck, 1816, p. 478; Risso, 1826, p. 305; Blainville, 1827, p. 392; 1816���1830 Pl. 49, fig. 1; 1830, p. 108;1834, p.118; delle Chiaje, 1829, p. 3, 29; Lesson, 1843, p. 490; Gegenbaur, 1853, p. 42, Pl. 18, figs. 5���10; Keferstein & Ehlers, 1861, p. 32, Pl. II, fig. 18���25; L. Agassiz, 1862, p. 370; Vogt, 1862a,b fig.3; Claus, 1863, Pl. 47, fig. 20b; Spagnolini, 1869, p. 641; 1870, p. 39; Fewkes, 1879, p. 292, Pl. 2; 1883, p. 79; Bedot, 1881, p. 122; Carus, 1885, p. 46; Haeckel, 1888b, p. 329; Chun, 1897a, p. 79, Pl. V, figs. 1���8; 1897b, p. 82, fig. 17, fig. 26a, 27; Schneider, 1898, p. 170; Lo Bianco, 1899, p. 463 Richter, 1907, p. 559, text-figs. 1-6, Pl. 27, figs. 1-13; Kawamura, 1910a, p. 447, Pl. XIV, fig. 3; 1954, p. 118; Lens and van Riemsdijk, 1908, p. 100, Pl. 18, figs. 141-145; Bigelow, 1911b, p. 319; 1915, fig. 81; 1918, p. 427; Leloup, 1955a, p. 17; Bigelow & Sears, 1937, p. 65, fig. 50; Totton, 1965, p. 41, Pl. I, fig. 2, Pl. II-IV; Patriti, 1970, p. 75, fig. 105; Daniel, 1974, p. 28, fig. 1, H-N; 1982, fig. 1 (4���6); 1985, p. 49, fig. 9; Stepanjants, 1977, p. 56, fig. 2,A,Б; Bonnemains & Carr��, 1991, p. 50, fig. 1; Pag��s, 1991, p. 306, fig. 8.3; Pag��s & Gili, 1992, p. 69, fig. 3; Pugh, 1999, p. 480, fig. 3.4; Gao et al, 2002, p.66, text-fig. 24; Dunn, et al., 2005, p. 922, fig. 5-8; Zhang, 2005, p. 16, figs. 1C, 12; Dunn & Wagner, 2006, p. 747, figs. 2, 7; Araujo, 2012, p.58. Rizophysa filiformis Brugui��re et a l. 1827, p. 139, Pl. 89, fig. 12; Epibulia filiformis Eschscholtz, 1829, p. 148. Rhizophysa sp. Fewkes 1881, p. 773, fig. 11 Rhizophysa gracilis Fewkes, 1882a, p. 269, Pl. VI, figs. 1���6; Cannophysa gracilis Haeckel, 1888a, p. 44. Cannophysa murrayana Haeckel, 1888a, p. 44; 1888b, p. 324, Pl. 24. Pneumophysa gegenbauri Haeckel, 1888b, p. 328. Cannophysa eysenhardti Mayer, 1894, p. 239, Pl. 3, figs. 1,2,4. Rhizophysa murrayana Chun, 1897a, p. 84; Mayer, 1900, p. 72.? Epibulia (Macrosoma) mertensii Brandt, 1835, p. 233? Rhizophysa mertensii Lesson, 1843, p. 492; Haeckel, 1888b, p. 329.? Pneumophysa mertsensi Haeckel, 1888b, p. 45? Cannophysa filiformis Mayer, 1894, p. 241, Pl. 3, fig. 3.? Rhizophysa clavigera Chun, 1897b, p. 104; Mayer, 1900, p. 72;? Rhizophysa (?) tricornuta Moser, 1925, p. 445, 533 textfig. 59.? Rhizophysa (?) tricarnuta Moser, 1925, Pl. 33 (legend). non Rhizophysa filiformis Huxley, 1859, p. 90, Pl. 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(Eds.), Diversidad Biol��gica en la Reserva de la Biosfera de Sian Ka���an, Quintana Roo, M��xico. Centro de Investigaciones de Quintana Roo /PTSC Universidad de Florida, M��xico, pp. 109���125 Gili, J.���M. (1986) Estudio sistematico y faunistico de los cnidarios de la Costa Catalina. Ph.D. thesis, Facultad de Ciencias, Universidad Aut��noma de Barcelona, 565 pp. Gillett, K. & Yaldwyn, J. (1969) Australian Seashores in Colour. Tuttle, Sydney, 112 pp. Goto, S. (1897) Die Entwickelung der Gonophoren bei Physalia maxima. Journal of the College of Science, Imperial University of Tokyo, 10, 175���191., pl. 15. Griffith, E. & Pidgeon, E. (1834) The Mollusca and Radiata arranged by the Baron Cuvier. Whittaker & Co., London, 601 pp., 15 pls. https://doi.org/10.5962/bhl.title.10154 Halstead, B.W. (1965) Poisonous and venomous marine animals of the world. I. Invertebrates, United States Government Printing Office, Washington, D.C., i���xxxv, 1���994. Hoeven, J. van der. (1856) Systematic arrangement of Acalephae pp. 108-126 in Handbook of Zoology, Vol. 1. University Press, Cambridge, 844 pp. + 16 pls. Hong, H. (1964) Studies on the medusae of the East China Sea. 1. Siphonophora. Contributions of the Fisheries College, Shanghai, 1, 111���130. [In Chinese] Iosilevskii, G. & Weihs, D. (2009) Hydrodynamics of sailing of the Portuguese man-of-war Physalia physalis. Journal of the Royal Society Interface, 6, 613���626. https://doi.org/10.1098/rsif.2008.0457 Janssen, J., Gibbs, R.H. & Pugh, P.R. (1989) Association of Caristius sp. (Pisces, Caristiidae) with a siphonophore, Bathyphysa conifera. Copeia 1989 No. 1, 198���201. https://doi.org/10.2307/1445624 Hulet, W.H., Belleme, J.L., Musil, G. & Lane, C.E. (1974) Ultrastructure of Physalia nematocysts. In: Humm, H.J. & Lane, C.E. (Eds,), Bioactive compounds from the sea. Marcel Dekker Inc., New York, pp. 99���114 Keferstein, W. & Ehlers, E. (1861) Beobachtungen ��ber die Siphonophoren von Neapel und Messina. Zoologische Beitr��ge Gesammelt im Winter 1859/ 60 in Neapel und Messina. Leipzig: Wilhelm Engelmann, 34 pp. https://doi.org/10.5962/bhl.title.4760 Kirkpatrick, P.A. & Pugh, P.R. (1984) Siphonophores and Velellids. Synopses of the British Fauna, 29, 154 pp. Koeppern, J.H. (1913) Scotia collections. Siphonophora of the Scottish National Antarctic Expedition. Proceedings of the Royal Physical Society of Edinburgh, 19, 16���21. Lamarck, J.B.P.A de M. de. (1840) Histoire naturelle des animaux sans vert��bres. 2nd Edition, 3 rd volume, Bailli��re, Paris, 770 pp. https://doi.org/10.5962/bhl.title.63986 La P��rouse, J.F. de G. (1793) Atlas du voyage de la Perouse. Paris. ???page numbers??? https://doi.org/10.5962/bhl.title.15961 Leloup, E. (1934) Siphonophores calycophorides de l���ocean Atlantique tropical et austral. Bulletin du Mus��e Royal d���Histoire Naturelle de Belgique, 10 (6), 87 pp. Leloup, E. 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(1759) Amoenitates Academicae; seu dissertationes variae Physicae, Medicae, Botanicae, antehac seorsim editae, nunc collectae et auctae, cum tabulis aenaenis. Volume 4. ???page numbers??? https://doi.org/10.5962/bhl.title.910 Linnaeus, C. von. (1760) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species; cum characteribus, differentiis, synonymis, locis. Vol. I. 10 th edition, 823 pp. https://doi.org/10.5962/bhl.title.7377 Linnaeus, C. von. (1767) Systema Naturae, per regna tria naturae, secundum classes, ordines, genera, species; cum characteribus, differentiis, synonymis, locis. 13 th Edition. Vol. I, Classis VI, 1364 pp. https://doi.org/10.5962/bhl.title. 68927 Linnaeus, C. von. (1772) Caroli a Linn�� Systema Naturae ex editione duodecima in epitomen redactum et praelectionibus acadamicic accommodatum a Iohanne Beckmanno. Vol. I. 250 pp. https://doi.org/10.5962/bhl.title. 36521 Lo Bianco, S. (1899) Notizie biologiche reguardanti specialmente il periodo di maturit�� sessuale degli animali del golfo di Napoli. Mitteilungen aus der Zoologischen Station zu Neapel, 13, 448���573., 13 pls. Marsh, L.M. & Slack-Smith, S. (1986) Sea Stingers, Western Australian Museum, Perth, 133 pp. Mayer, A.G. (1900) Some Medusae from the Tortugas, Florida. Bulletin of the Museum of Comparative Zoology at Harvard College, 37, 13���82., 46 pls. McGrady, J. (1857) Gymnophthalmata of Charleston Harbour. Proceedings of the Elliott Society, 1, 103���221., pls. 8���12. Migotto, A.E., Marques, A.C., Morandini, A.C. & da Silveira, F.L. (2002) Checklist of the Cnidaria Medusozoa of Brazil. Biota Neotropica, 2, 1���31. https://doi.org/10.1590/S1676-06032002000100010 Moser, F. (1924b) Ordnung: Siphonophora. I: K��kenthal, W. und Krumbach, W. Handbuch der Zoologie, Vol 1, 485���521. Pag��s, F. (1991) Ecologia y Systematica de los Cnidarios Planctonicos de la Corriente de Benguela (Atlantico Sudoriental). Ph. D. Thesis, University of Barcelona, 466 pp. Pag��s, F. & Gili, J-M. (1992) Siphonophores (Cnidaria, Hydrozoa) of the Benguela Current (southeastern Atlantic). Scientia Marina, 56 (Supl. 1), 65���112. Park, J.H. & Song J.-I. (2004) Two new records of Hydromedusae (Cnidaria: Hydrozoa) in Korea. Korean Journal of Systematic Biology, 20, 31���37. Patriti, G. (1970) Catalogue des cnidaires et ctenaires des cotes Atlantique Marocaines. Traveaux de l���Institut Scientifique Ch��rifien. S��rie Zoologie, No. 35, 1���149. Pickwell, G.V. (1967) Gas and bubble production by siphonophores. Department of the Navy. Naval Undersea Warfare Centre, NUWC TP8, 98 pp. Prieto, L., Macias, D., Peliz, A. & Ruiz, J. (2015) Portuguese Man-of-War (Physalia physalis) in the Mediterranean: A permanent invasions or a casual appearance? Scientific Reports, 5, 11545, 6 pp. https://doi.org/10.1038/srep11545 Pugh, P.R. (1999) Siphonophorae. In: Boltovskoy, D. (Ed.), South Atlantic Zooplankton, Backhuys Publishers, Leiden, pp. 467���511. Richter, W. (1907) Die entwicklung der Gonophoren einiger Siphonophoren. Zeitschrift f��r Wissenschaftliche Zoologie, 86, 557���618, pls. 27���29. Risso, A. (1826) Histoire naturelle des principales production de l���Europe M��ridional, Volume 5, Levrault, Paris, 403 pp., 10 pls. https://doi.org/10.5962/bhl.title. 48984 Southcott, R.V. (1982) Jellyfishes (Classes Scyphozoa and Hydrozoa). In: Shepherd, Published as part of Pugh, P. R., 2019, A history of the sub-order Cystonectae (Hydrozoa: Siphonophorae), pp. 1-91 in Zootaxa 4669 (1) on pages 87-91, DOI: 10.11646/zootaxa.4669.1.1, http://zenodo.org/record/3773373

Key to the species of the Family Rhizophysidae It should be noted that all four recognised rhizophysid species possess relatively large pneumatophores, compared with Physonectae species, and, uniquely have finger-like processes from the gas gland, called hypocystic villi, hanging down from the base of the pneumatophore. 1. Young gastrozooids with lateral wing-like structures, ptera...................................Genus Bathyphysa... 2. Young gastrozooids without lateral wing-like structures...................................... Genus Rhizophysa...3. 2. Tentacles without side branches, tentilla.................................................... Bathyphysa conifera Tentacles with side branches, tentilla....................................................... Bathyphysa sibogae 3. Tentacles with simple tubular tentilla................................................... Rhizophysa eysenhardtii Tentacles with three types of complex tentilla............................................... Rhizophysa filiformis, Published as part of Pugh, P. R., 2019, A history of the sub-order Cystonectae (Hydrozoa: Siphonophorae), pp. 1-91 in Zootaxa 4669 (1) on page 74, DOI: 10.11646/zootaxa.4669.1.1, http://zenodo.org/record/3773373

Physalia post-Haeckel At last, we can return to sanity! Or, to paraphrase Merrill (1949, referring to Rafinesque, as quoted by Endersby, 2009, p. 170): ���in taxonomy and nomenclature we would have been infinitely better off today had Haeckel never written or published anything appertaining to the subject���. It did not take long for Claus (1889a, b), the latter being an English translation of the former, to give a critical review of Haeckel���s 1888 contributions. Claus particularly took issue with Haeckel���s nomenclature for the parts and appendages of siphonophores. However, in actuality that probably had more to do with the translator than Haeckel himself, since the original text was written in German. Claus wrote a great deal in regard to Haeckel���s Medusome Theory, which he debunked very successfully and regarded it as ���fiction���! Indeed it attracted few followers. With regard to Haeckel���s classification of the cystonects, Claus (ibid. p. 196-7) stated: ���There are numerous novelties in connexion with the division of pre-existing genera into two or more, and, indeed, on the ground of trifling distinctions scarcely applicable as generic characters. As examples may be cited the division of Physalia into Physalia and Caravella and of Alopleota [sic] into Aloploeota [sic] and Arethusa as also the establishment of two subfamilies associated therewith; further the breaking up of Rhizophysa by its different species into the genera Aurophysa, Cannophysa, Linophysa, Nectophysa, Pneumophysa, and Rhizophysa, and the distinction of two subfamilies as Cannophysidae and Linophysidae upon differences which perhaps justify generic separation���. Indeed, Claus (ibid., pp 195-6) had earlier remarked: ���That Haeckel makes a very extensive, indeed almost unlimited, use of his skill in making new and suitable names, is certainly intelligible from the fact that he possesses this faculty in a very high degree and has developed it, by many years��� practice, into a speciality, in which at present no other naturalist can hope to equal him���! Chun (1897) was one of the last 19 th century reviewers of the family Physaliidae, He, like Huxley, drew attention to a couple of the main offenders, Tilesius, Olfers and Lesson, who had done so much to muddle the already confused taxonomy of the family; and, correctly, also added Louis Agassiz (1862) to this list but, being an adherent to Haeckel���s Medusome Theory, he was reticent to criticise that particular author. However, Chun clearly did not consider that Haeckel���s division of the family Physalidae into sub-families, dependent on the presence or absence of a crest, was valid. In fact he believed that there were only two valid Physalia species; one living in the Atlantic and the other in the Indo-Pacific. The main difference between the two species was the other Haeckelian character, in that the Atlantic form had numerous main tentacles, while the other had only one. So Chun, like Huxley (1859), adopted the specific name for the Indo-Pacific species, Physalia utriculus (La Martini��re) [but actually, Gmelin, see above]. For the Atlantic species Chun applied what he considered to be the principles of priority and gave it 52 The name P. pelagica was first used by Lamarck (1801, p. 356). Bosc (1802, p. 166) used the misspelt form P. pelasgica. Haeckel (1888a) attributed this name to Osbeck (1757), but that is pre-Linnaean. 53 Haeckel (1888b) used Tilesius (1813) as the authority although, as discussed above, it appears to be Tilesius (1810). 54 Haeckel (1888b) quotes Eschscholtz (1829) as the authority for Physalia utriculus, but Gmelin (1788, p. 3155) was the first to use the specific name for Medusa utriculus (see above). 55 Although Haeckel (1888b) quotes Lamarck as the authority, but in brackets he says ���P��ron... pl. xxix, fig.1. Lesueur & Petit (1807) should be the authority. 56 O.F. M��ller (1776) is the original authority for this species, under the name Medusa caravella. the name P. Arethusa, based on the description and name given by Browne (1756). However, that was contrary to the then emerging International Rule of Nomenclature, as Schneider (1898, p. 190) 57 pointed out when he said: ���Linnaeus established in the 10 th edition of Systema Naturae, the Atlantic Physaliid as Holothuria physalis. It follows, therefore, that as a species name physalis, but not arethusa or Caravella, is preferable over both. Chun (97, p. 87) violates the principles of priority (which he believed to be following), when he designated our form: Physalia arethusa Browne (see also the rules of nomenclature).��� Thus, after 140 years since it was first named by Linnaeus, Schneider finally established the name Physalia physalis by which the species is known today. Although Schneider (1898, p. 87) 58 also retained the name Physalia utriculus for the ���smaller and less luxuriously developed��� Indo-Pacific form, he commented (ibid.) 59: ���If there are no morphological differences in the construction of the individual appendages between the two forms, it seems to me that it should be understood that the Pacific form is only a variety of the Atlantic one.��� As noted above, it was clear that Schneider was not a friend of Chun, judging by their comments about each others��� work, some of which are definitely worth reproducing. For instance, Schneider (1898, p. 51) 60 began his review of the systematics of siphonophores by stating: ���For the present communication I am induced by the two Chun works of 1897 [a, b]. To my regret, I have found that Chun, when judging my second communication of 1896, ���Grundriss der Organisation der Siphonophoren,��� had very little objective preoccupation, and spoke so disparagingly about many points - without thereby substantiating his opposite opinion or my opinion against earlier statements to acknowledge the grounds for refutation - that in the interest of my own work I consider a detailed criticism of Chun���s recent communications to be useless.��� However, he was more forthright (ibid., p. 162) 61 when he stated: ���I cannot refrain from expressing my satisfaction that Chun now finally recognizes the extra-capsular attachment of the stinging tube in the young nematocyst. Chun still said ���92 on p. 156 against my statement of ���91: ���I regret having to contradict almost all of Schneider���s information.��� Perhaps the time will come when Chun will no longer have to contradict the information I have provided. Incidentally, a statement of the incorrectness of my findings is just as welcome as a confirmation, because I work in the interest of science. Only then must the inaccuracy be ���explained��� in detail; empty sarcastic phrases are not enough in scientific studies.��� Schneider (1898) gave a quite detailed account of the morphology of rhizophysids, particularly with regard to the structure of the pneumatophore and the gonodendra, and made many comments, not all positive, about Chun���s (1897) description, and, with regard to the giant cells of the pneumatophore, he stated (ibid., p. 167) 62: ���The interpretation that Chun presents for the giant cells appears completely untenable to me. He regards them as ���buffers,��� that ���prevent vigorous contractions of the stem and the pneumatocodon above it from rupturing the pneumatophore.��� If there is any element that would appear to be inappropriate to fulfil such a function it is precisely these giant cells, 57 Original text: ���Linn�� f��hrt in der 10. Auflage des Systema naturae die atlantische Physalie als Holothuria physalis auf.Als Speciesnamen ergiebt sich summit physalis, nicht aber arethusa oder caravella, die beide bevorzugt werden. Chun (97 p. 87) verst��sst gegen die Grunds��tze der Priorit��t (denen er doch zu folgen glaubt), wenn er unsre Form: Physalia arethusa Browne benennt (siehe dar��ber die Nomenclaturregeln).��� 58 Original text: ���kleiner und weniger ��ppig entwickelt��� 59 Original text: ���Falls nicht morphologische Differenzen im Bau der einzelnen Anh��nge zischen beiden Formen bekannt werden sollten, scheint mir die pacifische Form nur als Variet��t der atlantischen aufgefasst werden zu k��nnen.��� 60 Original text ���Zu der vorliegenden Mittheilung Averde ich durch die beiden Chun���schen Arbeiten von 1897 veranlasst. Ich habe zu meinem Bedauern constatieren m��ssen, dass Chun bei Beurtheilung meiner zweiten Mittheilung von 1896: ��Grundriss der Organisation der Siphonophoren ��sehr wenig objectiv vorgieng und sich ��ber manche Puncte derart abf��llig aussprach - ohne dabei seine entgegengesetzte Auffassung zu begr��nden oder meine gegen fr��here Angaben vorgebrachten Gr��nde einer Widerlegung zu w��rdigen - dass ich im Interesse meiner eigenen Arbeit eine ausf��hrliche Kritik der Chun���schen neueren Mittheilungen f��r n��thig erachte.��� 61 Original text ���Ich kann hier nicht unterlassen, meine Genugthuung dar��ber auszudr��cken, dass auch Chun nun endlich die extracapsul��re Anlage des Nesselschlauches in den jungen Nesselzellen anerkennt. Sagte doch Chun noch 92 auf p. 156 gegen��ber meinen Angaben von 91: ��Ich bedaure, fast s��mmtlichen Angaben von Schneider widersprechen zu m��ssen.�� Vielleicht wird auch die Zeit kommen, wo Chun den von mir 96 gemachten Angaben nicht mehr widersprechen zu m��ssen bedauert. ��brigens ist mir eine Darlegung der Unrichtigkeit meiner Befunde ebenso willkommen, wie eine Best��tigung, denn ich arbeite ja im Interesse der Wissenschaft. Nur muss die Unrichtigkeit dann auch eingehend ��dargelegt�� werden; leere sarcastische Phrasen gen��gen nun einmal bei wissenschaftlichen Untersuchungen nicht. 62 Original text: ���Ganz unhaltbar erscheint mir die Deutung, welche Chun f��r die Riesenzellen aufstellt. Er betrachtet sie als ��Puffer��, die ��bei energischen Contractionen des Stammes und des ihm aufsitzenden Luftschirmes ein Sprengen des Luftsackes verh��ten��. Wenn irgend welche Elemente dazu ungeeignet erscheinen m��ssten, so sind es gerade die Riesenzellen, die von ��usserst zart spongi��ser Beschaffenheit und ausserdem nur von einer sehr d��nnen St��tzlamelle umgeben sind. Mit demselben Recht k��nnte man Bl��tterteig als Puffer zwischen Eisenbahnwagen f��r geeignet halten.��� which are extremely delicate and of a sponge-like texture and, in addition, are surrounded only by a very thin supporting lamella. With the same reasoning one could argue that puff pastry is suitable as a buffer between railway carriages���. Schneider (1898) was possibly the first person to make a detailed study of the development of the ���cormidial��� units in Physalia. On his most developed specimens, he found 11 ���cormidial��� groups, five in the oral zone, and six in the main one. With regard to the gonodendra, he noted (ibid., p. 187) 63: ���The findings generally show that the genital groups are being detached in toto. In addition to the tentacle and its associated polyp you will find, if the genital group is no longer detectable, a short rounded stump, to which the genital group was originally attached. Since, as is well known, no medusa bud on Physalia itself reaches maturity, and rather here germ cells are completely lacking, thus the detached genital group seems to be the site of maturation, and in the future, in order to find more advanced stages, one must look for free-swimming groups when a swarm of Physalia occurs.��� As Schneider noted male gonophores on the gonodendra he examined, one presumes, therefore, that he considered both the male and female gonodendra to be detached before both the special swimming bells and the gonophores became mature. Steche (1910) made a much more detailed study of the order of development of the various ���cormidial��� groups, based largely on the same material that Chun had looked at. He followed Schneider (1898) in designating the Atlantic species, Physalia physalis, but was adamant that it was distinct from P. utriculus, mainly on the more complex nature of the arrangement of the various zooids in the former, and that there were marked differences in the development of the young zooids. He reviewed the other studies on developing larvae by Huxley (1859), Chun (1887) and Schneider (1897). He criticised the description by the last of these authors on several counts. In one instance he considered that Schneider���s conclusion that the ampulla at the base of the tentacle was very similar to the basigaster of a physonect gastrozooids, since in contained innumerable numbers of developing nematocysts, and which Schneider considered to have separated itself from the gastrozooid in order to give greater freedom to the tentacle, as his findings gave no support to that idea although it is not quite clear what these findings were. He also noted that the ���cormidial��� groups did not arise in a linear order and, thus, (ibid., p. 369) 64 he concluded: ���This proves that at an early stage germinative material must be distributed over the entire stem zone, which is then activated one after another���. Bigelow (1911) followed Steche in recognising the two species, Physalia physalis in the Atlantic, and P. utriculus in the Indo-Pacific. He pointed out that although the Pacific species, with its single main tentacle, resembled the immature stage of the Atlantic species; nevertheless of all the specimens from the Pacific that he, and previous authors, had examined none possessed more than one main tentacle. However, a year earlier Kawamura (1910), in a somewhat obscure paper in Japanese, had already shown that of his specimens from near Izu Oshima, Japan, two had two major tentacles while one had four. Kawamura, thus, considered utriculus to be a variety of P. physalis. Moser (1925), having studied a large collection of physaliid specimens from the Gauss and other expeditions, came to the same conclusion in that the Atlantic and Indo-Pacific forms were merely varieties of one species, which would thus be called Physalia physalis. Nevertheless, she still recognised the possibility of two varieties that she referred to as the Atlantic arethusa and the Indo-Pacific utriculus. The final nail in the coffin for the idea that there were two species of Physalia should have come from Okada (1932, 1935) who examined a large number of Physalia specimens from Japan with a pneumatophore ranging in length from 2 to 100 mm. He also made very detailed studies of the order of origin of the various zooids and, helpfully, used the same system of annotations as Steche (1910) had used. Unlike Steche, however, he finally concluded from his studies (ibid p. 19) 65: ���It is wrong, as was believed until now, that the Indo-Pacific Ocean Physalia has only one main tentacle, and that is because we had never observed a fully developed Physalia. The specimen, whose pneumatophore reached 95 to 100 mm long, has as many long and major tentacles as the species living in the Atlantic Ocean.��� 63 Original text ���Die Befunde lehren ganz allgemein, dass die Genitalgruppen in toto abgestossen werden. Neben dem Fang- faden und seinem zugeh��rigen Polypen findet man - falls die Genitalgruppe nicht mehr nachweisbar ist - einen kurzen abgerundeten Stummel, an dem urspr��nglich die Genitalgruppe festsass. Da nun, wie bekannt, keine Medusenknospe an der Physalia selbst zur Reife gelangt, alle vielmehr hier der Keimzellen vollst��ndig entbehren, so scheint die abgel��ste Genital- gruppe die Reifungsst��tte zu sein, und man wird k��nftig, um weiter vorgeschrittene Stadien zu finden, bei Auftreten eines Physalia-Schwarmes nach frei schwimmenden Gruppen suchen m��ssen.��� 64 Original text ���Dies beweist, dass schon fr��hzeitig Keimmaterial ��ber die ganze Stammzone verteilt sein muss, das nachein- ander aktiviert wird.��� 65 Original text: ���C���est �� tort que l���on a cru jusqu����� pr��sent que la Physalie de l���Oc��an Indo-Pacifique n���avait qu���un tentacule principal, et cela parce que l���on n���avait jamais observ�� la Physalie en parfait d��veloppement. Le sp��cimen dont le pneu- matozo��de atteint 95 �� 100 mm. de long, poss��de de longs et nombreux tentacules principaux, comme l���esp��ce vivant dans l���Oc��an atlantique.��� Totton (1960, p. 304), who made very detailed studies of the budding patterns of the ���cormidial��� groups, was clearly not particularly impressed with the studies on this subject, made by earlier authors when he stated: ���The methods employed had to be purely exploratory to begin with, because previous accounts of the arrangement of the groups of buds in Physalia were too superficial and illustrated either not at all (Schneider, 1896) or so inadequately (Steche, 1910; Okada, 1932, 1935) that they could not be checked and built upon. In fact it was necessary to start de novo ���. However, he did commend Steche (1907) for possibly being the first person to recognize that the gonophores, usually assumed to be always male, while the special nectophores were assumed to be the female, could also be female; and that all gonophores on a single gonodendron were of the same sex. Was this the first recognition that cystonects were dioecious? It is not clear. Totton (1960) also was adamant that there was only one Physalia species, namely P. physalis. In fact, all major reviewers since Bigelow (1911) have recognised only the one species. However, there are still a few exceptions to be found in the literature. Collins (2002) used material from an Atlantic (as P. physalis) and a Pacific (as P. utriculus) specimen in his analyses of the molecular phylogeny of the Medusozoa. In one instance (ibid. Fig.1) he found that that the ���Bootstrap indices under maximum parsimony and minimum evolution criteria��� for distinguishing the species were both 100; indicating that there was a high degree of likelihood that those two specimens were more closely related to each other than to anything else included in the analysis. However, although the likelihood is high, one cannot use these data to prove that one is dealing with a single species (Dr Casey Dunn, personal communication). As Dr Dunn has also commented that the more recent papers that have suggested that there is more than one Physalia species have largely been peripheral to the core siphonophore literature, as for more than 70 years siphonophore systematists have agreed that there is just one. For instance, at least two teams of medics have suggested ways to distinguish more than one species of Physalia. Fenner et al. (1993) suggested that the response to the experimental exposure of Physalia tentacles, from diff, Published as part of Pugh, P. R., 2019, A history of the sub-order Cystonectae (Hydrozoa: Siphonophorae), pp. 1-91 in Zootaxa 4669 (1) on pages 54-58, DOI: 10.11646/zootaxa.4669.1.1, http://zenodo.org/record/3773373, {"references":["Merrill, E. D. (1949) Index Rafinesquianum: the plant names published by C. S. Rafinesque with reductions, and a consideration of his methods, objectives and attainments. Arnold Arborteum of Harvard University, 296 pp.","Endersby, J. (2009) ' The vagaries of a Rafinesque': imagining and classifying American nature. Studies in History and Philosophy of Biological and Biomedical Sciences, 40, 168 - 178. http: // dx. doi. org / 10.1016 / j. shpsc. 2009.06.009","Claus, C. (1889 a) Zur Beurtheilung des Organismsus der Siphonophoren und deren phylogenetischer Ableitung. Eine Kritik von E. Haeckel's sog. Medusom-Theorie. 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Pl. 23, figs. 1 - 6.","Eschscholtz, F. (1829) System der Acalephen. 1 - 190, 16 pls. Berlin, 539 pp. https: // doi. org / 10.5962 / bhl. title. 10139","Gmelin, J. F. (1788) Caroli a Linne, Systema naturae per Regna Tria Naturae. Vol. I, Part VI. pp. 3021 - 3910. https: // doi. org / 10.5962 / bhl. title. 125970","Lesueur, C. A. & Petit, N. de (1807) Voyage de decouvertes aus terres australes. Atlas, 40 pls. Imprimerie Imperiale, Paris. https: // doi. org / 10.5962 / bhl. title. 44096","Muller, O. F. (1776) Beschreibung zweier Medusen. Beschaftigungen der Berlinischen Gesellschaft Naturforschender Freunde, 2, 290 - 297, pl. 9.","Browne, P. (1756) The civil and natural history of Jamaica. Osborne & Shipton, London, viii + 533 pp, 50 Pls. https: // doi. org / 10.5962 / bhl. title. 10826","Schneider, K. C. (1898) Mittheilungen uber Siphonophoren. III. Systematische und andere Bemerkungen. Zoologischer Anzeiger, 21, 51 - 53, 73 - 93, 114 - 133, 153 - 173, 185 - 200.","Steche, O. (1910) Das Knospungsgesetz und der Bau der Anhangsgruppen von Physalia. Festschrift. zum sechzigsten Geburtstag Richard Hertwigs, Jena. Vol. 2, 357 - 372, 10 Plates.","Chun, C. (1887) Zur Morphologie der Siphonophoren. 2. Ueber die postembryonale Entwickelung von Physalia. Zoologische Anzeiger, 10, 557 - 561, 574 - 577.","Bigelow, H. B. (1911) The Siphonophorae. Memoirs of the Museum of Comparative Zoology at Harvard College, 38, 171 - 402.","Kawamura, T. (1910) \" Bozunira \" and \" Katsuwo no Eboshi \" Rhizophysa and Physalia. Dobutsugaku zasshi Tokyo, 22, 445 - 454, 1 pl. [In Japanese]","Moser, F. (1925) Die Siphonophoren der Deutschen Sudpolar-Expedition, 1901 - 03. Deutsche Sudpolar-Expedition, 17 (zool 9), 1 - 541.","Okada, K. (1932) Developpement post embryonnaire de la Physalia pacifique. Memoirs of the College of Sciences, Kyoto Imperial University, B 8, 1 - 25, 1 pl.","Okada, K. (1935) Les jeunes physalis. Note supplementaire sur le development postembryonnaire de la physalie. Memoirs of the College of Science, Kyoto Imperial University Series B, 10 (5), 407 - 410.","Totton, A. K. (1960) Studies on Physalia physalis (L.). Part 1. Natural History and morphology. Discovery Reports, 30, 301 - 368.","Schneider, K. C. (1896) Mittheilungen uber Siphonophoren. II. Grundriss der organisation der Siphonophoren. (Jena) Zoologischen Jahrbucher. Abteilung fur Anatomie und Ontogenie der Tiere Abteilung fur Anatomie und Ontogenie der Tiere, 9, 571 - 664.","Steche, O. (1907) Die Genitalanlagen der Rhizophysalien. Zeitschrift fur Wissenschaftliche Zoologie, 86, 134 - 171.","Collins, A. G. (2002) Phylogeny of Medusozoa and the evolution of cnidarian life cycles. Journal of Evolutionary Biology 15, 418 - 432. http: // dx. doi. org / 10.1046 / j. 1420 - 9101.2002.00403. x","Fenner, P. J., Williamson, J. A., Burnett, J. W. & Rifkin, J. (1993) First aid treatment of jellyfish stings in Australia. Response to a newly differentiated species. Medical Journal of Australia, 158, 498 - 501.","Yanagihara, A. A., Kuroiwa, J. M. Y., Oliver, L. M., & Kunkel, D. D. (2002) Ultrastructure of nematocysts from the fishing tentacle of the Hawaiian bluebottle, Physalia utriculus (Cnidaria, Hydrozoa, Siphonophora). Hydrobiologia, 489, 139 - 150. https: // doi. org / 10.1023 / A: 1023272519668","Weill, R. (1934) Contribution a l'etude des Cnidaires et de leurs nematocystes. II. Valeur taxonomique du cnidome. Traveaux de la Station Zoologique de Wimereux, 11, 349 - 701.","Werner, B. 1965. Die Nesselkapseln der Cnidaria, mit besonderer Beruchsichtigung der Hydroida. 1. Klassifikation und Bedeutung fur die Systematik und Evolution. Helgolander Wissenschaftliche Meeresuntersuchungen, 12, 1 - 39.","Mariscal, R. N. (1974) Nematocysts. In: Muscatine, L. & Lenhoff, H. M. (Eds), Coelenterate Biology. Reviews and new Perspectives. Academic Press, pp. 129 - 178.","Ostman, C. (2000) A guideline to nematocyst nomenclature and classification, and some notes on the systematic value of nematocysts. Scientia Marina, 64 (Supl. 1) 31 - 46. https: // doi. org / 10.3989 / scimar. 2000.64 s 131","Pontin, D. R. (2009) Factors affecting the occurrence of stinging jellyfish (Physalia spp.) at New Zealand beaches. Ph. D. Thesis, Lincoln University, Canterbury, New Zealand. xiv + 129 pp.","Pontin, D. R. & Cruickshank, R. H. (2012) Molecular phylogenetics of the genus Physalia (Cnidaria: Siphonophora) in New Zealand coastal waters reveals cryptic diversity. Hydrobiologia, 686, 91 - 105. http: // dx. doi. org / 10.1007 / s 10750 - 011 - 0994 - 8","Gershwin, L-A., Zeidler, W. & Davie, P. J. F. (2010) Medusae (Cnidaria) of Moreton Bay, Queensland, Australia. Memoirs of the Queensland Museum, 54 (3), 47 - 108.","Bardi, J. & Marques, A. C. (2007) Taxonomic redescription of the Portuguese man-of-war, Physalia physalis (Cnidaria, Hydrozoa, Siphonophorae, Cystonectae) from Brazil. Iheringia, Serie Zoologia, 97, 425 - 433. https: // doi. org / 10.1590 / S 0073 - 47212007000400011","Munro, C., Siebert, S, Zapata, F., Howison, M., Damian-Serrano, A., Church, S. H., Goetz, F. E., Pugh, P. R., Haddock, S. H. D. & Dunn, C. W. (2018) Improved Phylogenetic Resolution within Siphonophora (Cnidaria) with implications for trait evolution. bioRΧ iv, 27 pp. https: // doi. org / 10.1016 / j. ympev. 2018.06.030"]}

The siphonophore sub-order Cystonectae presently comprises just five species in three genera and two families, and includes Physalia physalis, the Portuguese Man O’War. Despite the smallness of the group its history has been very chequered, particularly for P. physalis, which has been described under more than fifty different names. Haeckel (1888) was one of the worst offenders regarding the description of questionable species, but even Totton (1965) was uncertain as to the validity of some previously described cystonect species. Herein, an attempt has been made to review the history of the sub-order Cystonectae and to clarify its taxonomy. A list of synonyms for each recognised cystonect species is given in an appendix.

Bathyphysa conifera (Studer, 1878) Rhizophysa conifera Studer, 1878a, p. 4, Pl. 1, figs, 2, 4���7, Pl. 2, figs. 13���18; 1878b, p. 89; Lens van Riemsdijk, 1908, p. 106; Leloup, 1936, p. 19. Bathyphysa abyssorum Studer, 1878a, p. 21, Pl. 3; Haeckel, 1888b, p. 248; Chun 1897a, p. 104; Schneider, 1898, p. 171; Lens & van Riemsdijk, 1908, p. 105; Bigelow, 1911, p. 321, 351; Leloup, 1936, p. 19, Pl. I, fig. 7���8; Carr�� & Carr��, 1995, p. 561. Pterophysa grandis Fewkes, 1886, p. 969, Pl. 10, figs. 1-3; 1889, p. 515 Chun, 1897, p. 104; Lens & van Riemsdijk, 1908, p. 107, Pl. 19, Pl. 34, figs. 167���170; Bigelow, 1911, p. 321; Leloup, 1936, p. 20 Bathyphysa grandis Haeckel, 1888b, p. 249. Linophysa conifera Haeckel, 1888a, p. 45; 1888b, p. 326. Bathyphysa grimaldii Bedot, 1893, p. 4, Pl. 1; 1904, p. 14, Pls. 3���4; Chun, 1897, p. 104; Bigelow, 1911, p. 321, 351 Leloup, 1936, p. 21, Pl. I, fig. 1, 3���4; Kawamura, 1947, p. 95, figs. 1���5; 1954, p. 120, text-fig. 2, Pl. V, fig. 1���, Pl. VI, fig. 1���2, Pl. VII, fig. 1���7; Patriti, 1970, p. 75, fig. 106. Pterophysa abyssorum Schneider, 1898, p. 171. Pterophysa conifera Schneider, 1898, p. 171; Lens & van Riemsdijk, 1908, p. 106; Bigelow, 1911, p. 321. Pterophysa grimaldii Lens & van Riemsdijk, 1908, p. 107 Pterophysa (Bathyphysa) studeri Lens & van Riemsdijk, 1908, p. 111, Pl. 20, fig. 149, Pl. 22, figs. 153���155, 157���159, Pl. 23, fig. 165, Pl. 25, fig. 171; Bigelow, 1911, p. 321, 351; Leloup, 1936, p. 21. Bathyphysa conifera Leloup, 1936, p. 23, Pl. I, fig. 2, 5���6, Pl. II; 1955a, p. 14, fig. 4���6, 1955b, p. 17; Totton, 1965; p. 43, Pl. V, fig.3, Pl. VI, fig. 1���4; Janssen et al 1989, p. 198; Jones & Pugh, 2016, p. 1. Bathyphysa sibogae Lens & van Riemsdijk, 1908 Bathyphysa sibogae Lens & van Riemsdijk, 1908, p. 114, Pl. 20, fig. 148. Pl. 23, figs. 160���164, Pl. 24; Bigelow, 1911, p. 321, 351; Leloup, 1936, p. 22, Pl. I, fig. 9; Totton, 1965, p. 43, Pl. V, fig. 2; Biggs & Harbison, 1976, p. 14, fig. 1���4; Dunn & Wagner, 2006, p. 745, fig. 3; Physalia physalis Linn��, 1758 Holothuria physalis Linnaeus, 1758, p. 657; 1759, p. 254, Pl. III, fig. 6; 1760, p. 657; 1767, p. 1090; 1772, p. 198; Osbeck, 1765, p. 371 (non Plate 12 legend); 1771, p. 105, Pl. 12, fig. 1; Hjortberg, 1769, p. 226, Pl. VII (partim); Tor��n, 1771, p. 166, 262; Gmelin, 1788, p. 3139; Voigt, 1806, p. 507; Blumenbach, 1814, p. 443; Bennett, 1834, p. 6; Griffith & Pidgeon, 1834, p. 490, Zoophytes pl. 5, fig. 1; Chun, 1897a, p. 86;: Burmeister, 1885���1891, p. 2. Holothuria velificans Osbeck, 1765, Pl. 12, fig. 1; 1771 (Part 1, Pl. 12, fig. 1; Part 2, pp, 74; Tor��n, 1865, p. 166, 263; De Haan, 1827, p. 496; Chun, 1897a, p. 86. Medusa caravella O . F. M��ller, 1776, p. 190, Pl. 2, fig. 2; Gmelin, 1788, p. 3156; de Fr��menville, 1824a, p. 44; 1824b, p. 321 [both citing Linn�� as the authority]; De Haan, 1827, p. 500; Spagnolini, 1869, p. 642; 1870, p. 40. Meduse La Martini��re, 1787, p. 365, Pl. 2, figs. 13���14; Medusa utriculus Gmelin, 1788, p. 3155; La Perouse, 1797, atlas, Pl. 20, figs. 13���14; Blainville, 1830, p. 102; 1834, p. 113; Lamarck, 1840, p. 94; Fenner et al., p. 500, fig. 1; Gershwin et al, 2010, p. 79, Fig. 4 A���C; Chun, 1897a, p. 86. Physsophora physalis Modeer, 1789, p. 285, Pl. 10, figs. 1-2. Physalia pelagica Lamarck, 1801, p. 356; 1816, p.480; 1840, p. 92; Tilesius, 1810, p. 184; Eysenhardt, 1821, p. 421, Pl. 35, fig. 2; de Fr��minville, 1824a, p. 42; 1824b, pp. 320; de Haan, 1827, p. 496; Deslonchamps, 1828, p.468; Eschscholtz, 1829, p. 162; Olfers, 1832, p. 38; Bennett, 1837, p.43; 1860, p. 5; Lesson, 1843, p. 545; Thompson, 1844, p. 281; Huxley, 1859, p. 100; Chun, 1887, p. 557; 1897a, p. 87; 1897b, p. 8; Haeckel, 1888b, p. 351; Burmeister, 1885-91, p. 13; Mayer, 1900, p. 73. Physalia pelasgica Bosc, 1802, p. 159, Pl. 19, figs. 1-2; 1830, p. 198, Pl. 16, figs. 1-2; Bory de St Vincent, 1804, Vol III, p. 288, Pl. 54, fig. 1 (Atlas); Blainville, 1830, p. 103; Bennett, 1834, p.6;1837, p. 43; Chun, 1897a, p. 87. Physalia gigantea Bory de St. Vincent, 1804, Vol III, p. 289. Physalia Australis P��ron, 1807, p. 210; Lesson, 1826, Pl. 5, figs. 1-2; 1830, p. 38; Chun 1887, p. 558; 1897b, p. 8. Physalia megalista Lesueur & Petit, 1807, Pl. 29, fig. 1; De Haan, 1827, p. 496; Lamarck, 1840, p. 94; Olfers, 1832, p. 32. Physalis arethusa Tilesius, 1810, p. 298; Eysenhardt, 1821, p. 420, Pl. 35, fig. 1; Blainville, 1826, p. 131; De Haan, 1827, p. 497; Chun, 1897a, p. 558. Physalis glauca Tilesius, 1810, p. 300; Blainville, 1826, p. 131; 1830, p. 103; 1834, p. 113; De Haan, 1827, p. 496. Physalis pelagica Tilesius, 1810, p. 302; Lamarck, 1816, p. 480; Blainville, 1826, p. 132; 1834, p. 113; Lesson, 1830, p. 41. Physalis lamartinieri Tilesius, 1810, p. 309; Eschscholtz, 1825, p. 744; Blainville, 1826, p. 132; 1830, p. 103; 1834, p. 113 De Haan, 1827, p. 496; Olfers, 1832, p. 36. Physalis cornuta Tilesius, 1810, p. 314; Blainville, 1826, p. 132; 1834, p. 113; De Haan, 1827, p. 496. Physalis osbeckii Tilesius, 1810, p. 314; De Haan, 1827, p. 496. Physalia afer Tilesius, 1810, p. 314, Pl. 1, figs. 14, 16; De Haan, 1827, p. 496. Arethusa caravella Oken, 1815, p. 128. Physalis pelagica Lamarck, 1816, p. 480. Physalis tuberculosa Lamarck, 1816, p. 480; 1840, p. 93; Lesson, 1826, Pl. 5, fig. 3; 1830, p. 40; Blainville, 1834, p. 113; Chun, 1887, p. 558; 1897b, p. 8. Physalis megalista Lamarck, 1816, p. 481. Physalis elongata Lamarck, 1816, p. 481. Physalia arethus a Eysenhardt, 1821, p. 420, Pl. 35, fig. 1; Eschscholtz, 1825, p. 744; De Haan, 1827, p. 496, Blainville, 1830, p. 103; Lesson, 1830, p. 41; Olfers, 1832, p. 3, Pl. I, II; van der Hoeven, 1856, p. 112, L. Agassiz, 1862, pp. 335, 367, Pl. 35; A. Agassiz, 1865, p. 215, fig. 351; Fewkes, 1880, p. 139; 1883, p. 79; 1889, p. 518; Burmeister, 1885���91, p. 13: Chun, 1882b, p. 55; 1887, p. 557; 1897a, p. 89; Vanh��ffen, 1906, p. 36, figs. 55-57; Koeppern, 1913, p. 18, figs. 1-3. Physalia lamartinieri Eysenhardt, 1821, p. 421; De Haan, 1827, p. 496; Blainville, 1830, p. 113; Lesson, 1830, p. 41. Physalia osbeckii Eysenhardt, 1821, p. 421; De Haan, 1827, p. 496; Lesson, 1830, p. 41. Physalia Laperouse Eysenhardt, 1821, p. 421; Non Bory de St. Vincent, 1804. Physalia producta Olfers, 1822, pp. 347-356, Plate I; 1832, p. 32; De Haan, 1827, p. 498. Physalia obverse van Hasselt, 1823, p. 1414; 1824, p. 118; Olfers, 1832, p. 177. Physaliae Eichwald, 1824, p. 453, Pl. XV, figs. 1���7. Holothuria Thalia de Fr��menville, 1824a, p. 43; 1824b. p. 321. Physalia Thalia de Fr��menville, 1824a, p. 44; 1824b, p. 322. Physalia cristallina de Fr��menville, 1824a, p. 44; 1824b, p. 322. Physalia hyalina de Fr��menville, 1824a, p. 44; 1824b, 323. Physalia gaimardi Blainville, 1826, p. 132; 1830, p. 103; 1834, p. 113; De Haan, 1827, p. 498 Physalia atlantica Lesson, 1826, Pl. 4, figs. 3���4; 1830, p. 36; Griffith & Pidgeon, 1834, Pl. 5, fig. 1; Burmeister, 1885���91, p. 13; Chun, 1887, p. 557; 1897b, p. 8. Physalia antarctica Lesson, 1826, Pl. 5, fig. 2; 1830, p.39; Burmeister, 1885���91, p. 13. Physalia azoricum Lesson, 1826, Pl. 5, fig. 4; 1830, p. 42; 1843, p. 555. Physalia Boiei De Haan, 1827, p. 502. Thalia Brugi��re et al. 1827, p. 159, P. 89, fig. 1. Physalis cystisoma Lesson, 1828, Pl. after p. 466; 1830, p. 39; 1843, p. 557. Physalia elongata Lessons, 1828, p. 468; Lamarck, 1840, p. 9. Physalia caravella Eschscholtz, 1829, p. 160, Pl. 14, fig. 1; Gegenbaur, 1853, p. 340; Huxley, 1859, p. 100; Spagnolioni, 1869, p. 642; Carus, 1885, p. 49; Chun, 1887, p. 557; 1888a, p. 1173; 1888b, p. 245;1897b, p. 8; Bigelow, 1891, p. 90. Physalia utriculus Eschscholtz, 1829, p. 163, Pl. 14, fig.2; Blainville, 1834, p. 113; Lamarck, 1840, p. 94; Lesson; 1843, p. 557; Leuckart, 1851, p.190, Pl. 6, figs. 1-6; Huxley, 1859, p. 101, Pl. 10, Pl. 12, fig. 12; Haeckel, 1888b, p. 351; Burmeister, 1885���91, p. 13; Chun 1887, p. 558; 1897a, p. 86; 1897b, p. 8; Agassiz & Mayer, 1902, p. 169, Pl. 10, figs. 43-44; Browne, 1904, p. 744; Lens & van Riemsdijk, 1908, p. 118, Pl. 24, figs. 175���175; Steche, 1910, p. 361,figs.1���8; Bigelow, 1911, p.321; 1919, p. 347; Browne, 1926, p. 84; Kawamura, 1954, p. 120; Barnes, 1963, p. 7, figs. 1-6; Halstead, 1965, p. 300; Williamson et al., 1996, p. 137; Alam et al, 2002, p. 9; Collins, 2002, p. 420; Yanagihara et al, 2002, p. 139. Physalia eschholtzi Olfers, 1832, p. 37. Physalus arethusa Blainville, 1834, p. 113. Physalia (Salacia) megalis ta Brandt, 1835, p. 237 Physalia (Salacia) pelagica Brandt, 1835, p. 237 Physalia (Alophota) olfersii Brandt, 1835, p. 237; Lesson, 1843, p. 559. Physalia olfersii Quatrefages, 1854, p. 109, Pl. 3���4. Physalia ambigua Brandt, 1835, p. 237. Physalia tuberculosa Lamarck, 1840, p. 93 Physalia megalista Lamarck, 1840, p. 94. Physalia elongata Lamarck, p. 94. Physalia aurigera McCrady, 1857, p. 74; Fewkes, 1880, p. 144; Alophota olfersii Haeckel, 1888a, p. 46 Alophota giltschiana Haeckel, 1888a, p. 46; 1888b, p. 348, Pl. 26, figs. 1-3; Chun, 1897a, p. 87. Arethusa challengeri Haeckel, 1888a, p. 46; 1888b, p. 349, Pl. 26, figs. 4-8; Chun, 1897a, p. 87. Alophota mertensii Haeckel, 1888b, p. 348. Arethusa thalia Haeckel, 1888b, p. 349. Caravella maxima Haeckel, 1888b, p. 352. Caravella gigantea Haeckel, 1888b, p. 352. Physalia physalis Schneider, 1898, p. 190; Richter, 1907, p. 571, text-figs. 7-12, Pl. 27, figs. 14���18, Pl. 28, figs. 19���26; Steche, 1910, p. 358, figs. 9-10; Bigelow, 1911, p. 322; Moser 1924, fig. 510; 1925, p. 452; Leloup, 1934, p. 2; 1955a, p. 17; 1955b, p. 17; Woodcock, 1944, p. 195, fig.46; 1956, p. 253; Wilson, 1947, p. 139;, Pl. I-III; Totton, 1954, p. 15; 1960, p. 301; 1965a, p. 39, Pl. I, fig. 1; Fagetti, 1958, p. 189, fig.; Tregouboff & Rose, 1957, p. 349, pl. 78, figs. 1���5; Lenhoff & Schneiderman, 1959, p. 452, figs. 1���8; Mackie, 1960, p. 369, text-figs. 1���5, Pls. XXVI-XXVIII; 1965, p. 439, fig. 1;Totton, 1960. p. 303, text-figs. 1���31, Plates VII-XXV; 1965, p. 39, figs.1, 5, 6, Pl. I, fig. 1; Daniel & Daniel, 1963c, p. 189, fig. I, 11���13; Stepanjants, 1966, p. 1619, figs. 2,7; 1967, fig. 1Б, 12, 17, 28, 40-41, 54A; Pickwell, 1967, p. 9, figs. 2,7; Gillet & Yaldwyn, 1969, p. 34, textfigs, 18-19, Pls. 15���16; Patriti, 1970c, p. 76, fig. 107; Hulet et al., 1974, p. 99, fig. 1-8; Daniel, 1974, p. 22, Text-fig. 1A-G; 1982, fig. 1(1���3); 1985, p. 45, fig. 8a-e; Kirkpatrick & Pugh, 1986, p. 26, fig. 5; Southcott, 1982, p. 124, Pl. 13/5; Gili, 1986, p. 265, figs. 4.62k,n, 4.113i; Marsh & Slack-Smith, 1986, p. 13, figs. 9���10; Gasca, 1990a, p. 41, Pl. I, fig. 1; 1990b, p. 111, fig. 1; Bonnemains & Carr��, 1991, p. 39, fig. 9; Pag��s, 1991, p. 304, fig. 8.1; Fenner at al., 1993, o. 500, fig.2; Carr�� & Carr��, 1995, p. 562, fig. 184; Wrobel & Mills, 1998, p. 45, fig. 59; Pugh, 1999a, p. 479, fig. 3.1; Sutherland & Tibballs, 2001, p. 609, figs.26.13-14; Gao et a l, 2002, p. 64, text-fig. 23; Migotto et al 2002, p. 21, fig. 27; Bouillon et al, 2004, p. 209, fig. 121B���D; 2006, p. 441, Fig. 207A���C; Dunn et al, 2005, p.917, fig.1a; Zhang, 2005, p. 15, fig. 1B, 11; Araujo, 2006, p. 48, Pl. 1, fig. 1; 2012, p. 57; Bardi & Marques, 2007, p. 425, figs. 1���36; Iosilevskii & Weihs, 2009, p. 613, figs. 1,8; Prieto et al, 2015, p. 1; Araya et al, 2016, p. 731. Physalia physalis var. utriculus Kawamura, 1910, p. 449, text-figs. 4���6, Pl. XIV, figs.2-6; Dakin & Colefax, 1933, p. 198; 1954, p. 124; Hong, 1964, p. 117, Pl. IX, fig. 1; Tokioka, 1970, p. 183; 1973, p. 141; Park & Song, 2004, p. 34, fig. 2A-D. Physalis physalis Birsa et al., 2010, p. 429. Rhizophysa eysenhardtii Gegenbaur, 1859 Rhizophysa eysenhardtii Gegenbaur, 1859, p. 408, Pl. 32, figs. 46���49; Fewkes, 1883, p.82, Pl. 1, fig. 1; Chun, 1897, p. 83, Pl. V, fig. 9; Mayer, 1900, p. 72; Lens & van Riemsdijk, 1908, p. 103, Pl. 20, fig. 47, pl. 21, fig. 150, Pl. 24, fig. 172; Kawamura, 1910, p. 445, Pl. XIV, fig. 1���2; 1954, p. 118; Bigelow, 1911, p. 320; Leloup, 1934, p. 2; 1936, p. 15; 1956, p. 475; Daniel & Daniel, 1963, p. 190, fig. II, 1���2; Totton, 1965, p. 42, Pl. I, fig. 3, Pl. V, fig. 1, Pl. VII; Daniel, 1985, p. 53, fig. 10; Pag��s, 1991, p. 305, fig. 8.2; Pag��s & Gili, 1992, p. 68, fig. 2; Pugh, 1999, p. 480, fig. 3.3; Dunn et al, 2005, p. 922, fig.5���9. Nectophysa wyvillei Haeckel, 1888a, p. 45; 1888b, p. 327, Pl. 23; Agassiz & Mayer, 1902, p.169, Pl.8, fig. 36. Aurophysa ordinata Haeckel, 1888a, p. 44; 1888b, p. 323. Pneumophysa mertensii Haeckel, 1888a, p. 45; 1888b, p. 328. Rhizophysa mertensii Schneider, 1898, p. 170. ? Rizophysa planestoma Lesueur & Petit, 1807, Pl. 39, fig. 3; Eschscholtz, 1829, p. 147; Blainville, 1834, p. 118 [spelt planostoma]; Lamarck, 1840, p. 83; Lesson, 1843, p. 491. ? Rhizophysa planestoma Eschscholtz, 1829, p. 147; Blainville, 1834, p. 118; Lesson, 1843, p. 491. ? Rhizophysa filiformis Huxley, 1859, p.90, pl. 8, fig. 13���20. ? Rhizophysa inermis Studer, 1878a, p.13, Pl.1, figs. 3, 8���10. ? Aurophysa inermis Haeckel, 1888a, p.44; 1888b, p. 324., Published as part of Pugh, P. R., 2019, A history of the sub-order Cystonectae (Hydrozoa: Siphonophorae), pp. 1-91 in Zootaxa 4669 (1) on pages 84-87, DOI: 10.11646/zootaxa.4669.1.1, http://zenodo.org/record/3773373

Cnidarians are considered ancestral metazoans and, therefore, are important taxa for studying animal evolution. However, little is known about the group’s genome size (C value), which is an important parameter in whole-genome sequencing. To address this issue, we measured the C values of 27 cnidarian species from Japan, using flow cytometry, and found that they ranged from 0.26 to 3.56 pg. Excluding the results for Agalma elegans and Physalia physalis (order Siphonophorae), which had the highest C values among the species included in the present study, the C values for the cnidarians were 0.26–1.49 pg. In particular, we found that hydrozoans possessed relatively large and wide-ranging C values, indicating that evolution within the group involved considerable gains or losses of genomic content. Overall, the C values reported in the present study could be valuable for whole-genome sequencing, using next-generation sequencers, and for future research in cytogenetics.

Marine pleustonic species such as the hydrozoans Velella velella and Physalia physalis , are known to drift in the world's oceans driven by winds, currents and tides. Here we present the first chemotaxonomic characterization, based on the fatty acid (FA) profile, of these two charismatic oceanic species that thrive in the interface layer between air and the water column in adult stages. Moreover, we compared their FA profiles with those from other representative cnidarian orders (Rhizostomeae, Anthomedusae, Siphonophorae, Alcyonacea, Scleractinia, Helioporacea and Pennatulacea). Velella velella and P. physalis mainly differed in the presence of symbiotic dinoflagellates markers (18:3 n -6, 18:4 n -3 and 20:5 n -3 polyunsaturated FAs), present in higher percentage in the former, and bacterial markers (odd-numbered, branched and 18:1 n -7 FAs), which were more representative in the latter. When comparing these species' FA profiles with the ones of other cnidarians orders, the presence/absence of endosymbionts and of specific FAs (tetracosapentaenoic and tetracosahexaenoic acids) as well as the latitudinal habitats were the main drivers for the distinction between groups.

During summer 2017 an anomalous and unprecedented developing of warm water off the coast of Chile was presented, namely Coastal El Nino event. Coincidentally, a high amount of strandings of Portuguese Man-of-War (Physalia physalis) were present at that time. In this work we determine the latitudinal and seasonal distribution of the pleustonic siphonophore P. physalis in the southeastern Pacific, from January 2016 to December 2017. The northernmost and southernmost records for the southeastern Pacific were recorded in this study, from Arica (18 ∘ 26’S; 70 ∘ 18’W) to Quellon (43 ∘ 21’S; 74 ∘ 07’W) in Chile, respectively. During the surveyed period, the greatest strandings of this species were recorded both in the north (23 ∘ S) and south of Chile (36–38 ∘ S) during the austral summer months, coinciding when sea surface is relatively warmer. The number of strandings registered in 2017 was more than 4 times the number of strandings during 2016; this is 17339 colonies in 2017 and 3910 colonies in 2016. The association between the maximum observed strandings and the time of appearance of Coastal El Nino event during summer 2017 are also discussed. Our findings can be used as guidance in the monitoring of P. physalis, in order to prevent encounters and therefore possible accidents with humans in coastal waters and beaches by strandings.

9 pages, 5 figures, 3 tables, This study analysed the distribution and abundance of both polygastric (asexual) and gonophores of the calycophoran siphonophores Muggiaea atlantica and Muggiaea kochii in the Mondego estuary (Portugal), as well as the influence of local environment and prey availability on the observed patterns. Samples were collected in three different zones of the estuary, from December 2013 to November 2015. In general, both species nectophores and total gonophores followed a similar trend, with higher abundances during spring-summer periods and increased densities in the downstream areas of the estuary, where the marine influence is stronger. Inter-annual differences occurred mainly for M. atlantica nectophores, which showed higher abundances in 2015, particularly in the upstream areas. Both species displayed a shift in the annual peak abundance, from bimodal in 2014 to unimodal in 2015. Patterns observed were not only strongly related with an increase of the marine influence in the estuary, but also with the presence of zooplanktonic prey, such as cirripede cypris, calanoid nauplius, Poecilostomatoida, and copepodites of Cyclopoida. This study represents a step towards a better knowledge of the biology and ecology of siphonophores, The present work was supported by FCT (Portuguese Foundation for Science and Technology) through a PhD grant awarded to Mariaelena D'Ambrosio (SFRH/BD/91541/2012), the postdoctoral grants attributed to SC Marques (SFRH/BPD/110400/2015) and AL Primo (SFRH/BPD/91030/2012), the Investigator FCT program attributed to F Martinho (IF/01410/2012), the COMPARE Project (FCOMP-01-0124-FEDER- 020850) financed by POPH (Portuguese Operational Human Potential Program), QREN Portugal (Portuguese National Strategic Reference Framework), and MCTES. E Guerrero is part of the Marine Biodiversity Conservation Group (MEDRECOVER) 2014SGR-1297

Tottonophyes enigmatica sp. nov. Diagnosis. With two rounded, almost spherical, dissimilar nectophores with one slightly larger and slightly to the anterior of the other. Extensive nectosac and hydroecium occupying most of each nectophore. No flaps on lateral walls of hydroecium; no mouth plate on posterior nectophore; ascending and descending mantle canals present, with internal somatocyst forming a conical structure. No disjunct portion of the pedicular canal in either nectophore. Material examined. Four specimens of Tottonophyes enigmatica sp. nov. have been examined. All were collected by the MBARI ROVs, namely Tiburon (T), Ventana (V), or Doc Ricketts (DR), whose dive details are shown in Table 1. They were preserved in 5% buffered formalin in sea water. Unfortunately, only one of the nectophores from the DR0105 and DR0552 dives was preserved. A fifth specimen (T0398) was collected and photographed in the ship-borne laboratory, then part was frozen for molecular analyses. Five other specimens (unemboldened) in Table 1 have been identified from in situ frame grabs. The specimen from Tiburon Dive 897 has been designated as the holotype, and will be deposited at the Smithsonian National Museum of Natural History. The remainder will be deposited with the Peabody Museum of Natural History at Yale University. Description: An in situ frame grab from Doc Ricketts Dive 105 (Figure 5A) shows that the larger of the two nectophores of Tottonophyes enigmatica sp. nov. is only partially superimposed over the smaller one. This superimposition (i.e. positioning of one nectophore to the anterior of the other) is much greater than seen in prayomorphs (Figure 2A), where they are apposed, and much less than for other diphyomorphs, even the clausophyids (Figure 5B, C). However, unlike prayids, there are clear differences in the two nectophores of T. enigmatica sp. nov., and for ease of comparison with the diphyomorph species the nectophores will be referred to as anterior (larger) and posterior (smaller). The holotype specimen from Tiburon Dive 987 was chosen as the best preserved of all the specimens collected up until the time that this description was begun. Very recently a further specimen was collected during Doc Ricketts Dive 965. This proved to be the largest specimen so far collected, with the anterior nectophore measuring 13.5 mm in length and 9 mm in width, and the posterior one 11.5 by 7 mm. Nonetheless, the general structure of both nectophores showed no major variations from the norm, and there was no evidence there to suggest the unlikely eventuality that we were dealing with a different species. However, there appeared to be a minor difference in the special nectophores in the cormidia that is discussed below. Anterior nectophore: (Figures 6 & 7). The larger, anterior nectophore of the holotype specimen measured, in its preserved state, 7.6 mm in height, 7.6 mm in width and 4.6 mm in breadth. The proximal two-fifths of the nectophore was almost entirely occupied by the hydroecium (Figure 7), which was open at its lower end, but at its upper end its median wall curved over and ran to the proximal wall of the nectophore a short distance from its apex. The distal three-fifths of the nectophore were mainly occupied by the extensive nectosac; its ostial opening, which included a large velum, was directed obliquely toward the upper side of the nectophore, such that outer side of the nectophore was only about two-thirds the height of the inner or proximal side. All four radial canals arose from the short and direct internal pedicular canal. In its preserved state the upper part of the nectosac was more extensive laterally than in the mid-line, such that the upper radial canal ran through a median depression. However, in life this depression was less marked (Figure 6A). The upper and lower radial canals ran directly to the ostial ring canal. The lateral radial canals, however, followed a course somewhat similar to that found in Clausophyes species. From their origin, they extended across the nectosac, with only a very slight upward curve. Close to the outer side of the nectosac they curved through 90° and ran directly toward the ostium, before again bending through 90° and running back across the nectosac, with a slight upward curve as they approached the proximal side. A further curve through 90° again directed them toward the ostium, but before reaching it, they curved through c. 60° and then ran obliquely down to join with the ostial ring canal. FIGURE 7. Nectophores of Tottonophyes enigmatica sp. nov. A. anterior, and B posterior. h an . h pn: hydroecium of anterior and posterior nectophore, respectively; mca, mcd, ascending and descending mantle canal, respectively; ns: nectosac; pde, pdi, external and internal pedicular canal, respectively; soan, sopn: somatocyst of anterior and posterior nectophore, respectively. Scale bar 2 mm. As well as the short internal pedicular canal, which reached the nectosac approximately at the mid-height of the nectophore, there were two short median longitudinal canals running along the wall of the hydroecium: the ascending and descending mantle canals. The ascending canal was shorter than the descending one, and soon gave rise to the internal somatocyst, within the mesogloea. The somatocyst was a funnel-shaped, laterally compressed structure (Figures 6 & 7, soan). Its apex, which was covered with large globular cells, was expanded laterally so that, from this angle, the somatocyst looked like a mushroom (Figure 6B soan). In life, the apex of the somatocyst was deep brown in colour, while the remainder of the cone was suffused with a lighter tan colouration. For the DR0965–D 1 specimen the fluid within the somatocyst, particularly that of the anterior nectophore, in life, was a pinkish-red in colour. Posterior nectophore: The smaller posterior nectophore (Figures 6 & 7) measured, in its preserved state, 5.6 mm in height and 4.5 mm in width. Its breadth was not measured as the nectosac had collapsed and the whole nectophore narrowed. However, the posterior nectophore from Doc Ricketts Dive 105 remained in good condition and its dimensions were: height 7.6 mm, width 8.2 mm, breadth 6.2 mm. Thus, it was considerably larger than the posterior nectophore of the type specimen and, commensurately one would have expected the anterior nectophore to be considerably larger. However, as noted above, the anterior nectophore was unfortunately lost during collection. The hydroecium occupied less than a third of the total width of the nectophore (Figure 7) and was open throughout almost the entire height of the nectophore, but faded out shortly before reaching its lower edge. At its upper end, just above the swollen part of the internal somatocyst, the hydroecium formed a small median cavity. The remainder of the nectophore was almost entirely occupied by the nectosac, with its ostial opening and a slightly smaller velum directed obliquely toward the upper side of the nectophore. No mouth plate was present. The courses of the radial canals were similar to those of the anterior nectophore. In the preserved state, the upper canal likewise ran through a median furrow in the apical part of the nectosac, whose lateral wings apically came to a more or less pronounced point. This arrangement was also apparent in the living specimen. The lower canal ran directly down to the ostial ring canal. The lateral radial canals, after arising from the short internal pedicular canal, almost immediately curved basad and then after a short distance ran outwards across the nectosac. As they approached its outer wall they gradually curved to run basally again, before turning to run obliquely upward toward the proximal side. Then they curved basally again before turning distad and running obliquely down to the ostial ring canal. Both descending and ascending mantle canals were present (Figure 7 mcd, mca), although their relative lengths were the reverse of those in the anterior nectophore. The ascending mantle canal was much longer than that in the anterior nectophore and, below the median cavity in the hydroecium, it penetrated into the mesogloea and formed the internal somatocyst. This was relatively short and without lateral extensions (Figures 6 & 7 sopn). It also had a small brownish patch of cells at its distal end, while the remainder contained light-brown pigmentation, after preservation. Cormidium: The siphosomal gastrovascular lumen, like that of the somatocysts, contained a pinkish-red fluid. Much of the siphosome of the type specimen of Tottonophyes enigmatica sp. nov. was lost during collection and so only the very youngest immature cormidia remained (Figure 6). Fortunately, more of the siphosome was retained with the posterior nectophore collected during Doc Rickets Dive 105, and the terminal cormidia, although still immature, can be described. Photographs of the oldest one of these are shown in Figure 8. It consisted of a bract, a gastrozooid and tentacle, and what appears to be an asexual swimming bell, together with two small buds (Figure 8 b). The specimen from Doc Ricketts dive 552, received after the major part of the manuscript had been written, possessed a longer piece from the anterior end of the siphosome, despite the fact that the anterior nectophore was lost. Some of the cormidial groups appeared to be fully developed and as the swimming bells, at all stages of development, bore no trace whatsoever of a manubrium, then the presumption that they were special nectophores appeared to be correct. Bract: The largest bract Doc Ricketts Dive 552 specimen (Figure 9, left) measured 3 mm in length and 1.85 mm in width. It was a very simple, leaf-like structure, being concave on the lower side, convex on the outer. There was a small notch (Figure 10 A) toward the proximal end. The phyllocyst was reduced to a central, thickened part overlying the stem, and two hydroecial canals, which in the youngest bracts ended in slight swellings. In the younger bracts (Figure 10 B) the two hydroecial canals were of almost equal length, but as they enlarged they became distinctly asymmetrical. Special cormidial nectophore: The large medusoid (Figure 9, 10 C) structure measured about 2.5 mm in height and 2 mm in maximum width. It was a ridgeless, simple structure, largely occupied by the subumbrella cavity. All the radial canals were located laterally and originated from the short pedicular canal. One pair ran directly to the ring canal, while the other pair described broad curves, first upward then downward, before running directly to the ring canal. As none of these medusoids, at any stage in development, showed any signs of sexual products they were, as noted above, presumed to be special nectophores. For the more mature cormidial groups, as well as the large medusoid there were one or two small buds (Figure 8B) attached to the stem. These were presumed to be developing male and/or female gonophores but, in all cases, they were too small to be identified positively as such. As noted above, the special nectophores of the largest, recently collected specimen, from Doc Ricketts Dive 965, showed a slight difference from the norm in that the lateral radial canals did not follow the simple course as shown in Figure 10C. Instead their courses more closely resembled those of the lateral canals on the nectophores. On the other hand, the bracts showed no differences, with the two hydroecial canals and the thickened longitudinal one. As none of the other specimens had even partially developed cormidia, it is not, as yet, possible to say whether this difference is significant. Gastrozooid and tentacle: The small gastrozooids (Figure 11 A) measured about 12.5 mm in length and 7 mm in diameter, with the conspicuous basigaster occupying the proximal third of the whole. The tentilla (Figure 11 B) were of the typical calycophoran form with a C-shaped cnidoband, orange-red in colour, and a long terminal filament. The cnidoband bore two types of nematocyst. There were about 10 pairs of large nematocysts, measuring c. 95 x 20 µm, which appeared to be the microbasic mastigophores normally found for calycophorans; and several rows of smaller ones, measuring 28 x 5 µm, which were anisorhizas. The terminal filament contained desmonemes, 11 x 9 µm, and anisorhizas, but their exact arrangement was not investigated. Eudoxid stage ? Recently, a strange eudoxid (Figure 12) was found, as bycatch, in a suction sample collected during Doc Ricketts Dive 918 (17th December; 2016; 36°35.5’N, 122°25.57’W; water depth c. 2900 m) in Monterey Canyon, at a depth of 550 m. The eudoxid appeared to belong to a clausophyid species as there were two long hydroecial canals, extending down into the extensive neck shield of the bract. The young bracts of Tottonophyes enigmatica sp. nov. (see. Figures 9, 10) also showed these canals. The only non-clausophyid species whose eudoxid bracts show that character is Gilia reticulata (Totton) (see Pugh & Pagès, 1995) that, for the present, is placed in its own sub-family, Giliinae, within the family Diphyidae, until some phylogenetic analyses can prove otherwise. Unfortunately, although several attempts have been made, molecular analyses of G. reticulata have always failed. In its preserved state, the eudoxid bract measured 6.1 mm in length, of which the neck shield occupied about ¾rd. The headpiece was ridgeless and smooth, and was largely filled by the phyllocyst, which was globular in shape, measuring 1.35 mm in height, and 1.1 mm in width, with a narrow apical extension, 0.35 mm in length. The two hydroecial canals, which arose from the longitudinal canal, extended almost to the base of the neck shield. The longitudinal canal was well-defined and curved around the base of the phyllocyst, on the opposite side to the gonophore, and extended for a short distance beyond the point of origin of the hydroecial canals. As with the young cormidia, there appeared to be a special nectophore, 7 mm in length, because no sexual products were observed within its subumbrella cavity, which occupied almost the full length of the structure. The lateral radial canals arose from the internal pedicular canal at some distance below the apex of the nectophore. There was a distinct, rounded mouth plate. The deep hydroecium extended for almost the entire length of the zooid, with its lateral edges petering out on the mouth plate. The right-hand wing formed an extensive fold that overlapped the left-hand one for most of its length. Slightly above the ostial level, the right-hand fold was abruptly truncated, except on its outer margin where it protruded basally to form a distinct tooth-like structure The gastrozooid remained attached to the stem and at its base the tentacle was attached on one side while on the other were a few small buds, measuring up to 0.28 mm, that could be developing gonophores. Because of the presence of a pair of hydroecial canals, it would appear that this eudoxid belonged to a clausophyid species but, despite the similarities, we cannot be certain for the moment whether it belongs to Tottonophyes enigmatica sp.nov. However, given that the eudoxid stages, where present, of all clausophyid species are known, there remains a distinct possibility that it does. The eudoxid stages in clausophyid species vary considerably from the highly complex "fuseudoxid" of Crystallophyes (Pagès & Pugh, 2002) to, apparently, the complete absence of bracts, and hence eudoxids, in Clausophyes species. For the other genera, the eudoxid bract may be ridged (Chuniphyes and Heteropyramis) or smooth (Kephyes), but always the phyllocyst has an apical extension, and two distinct hydroecial canals. Distribution. Tottonophyes enigmatica sp. nov. is known from only a small area of the North East Pacific Ocean in the vicinity of Monterey Bay, California (see Table 1). Seven of the specimens, either collected or identified from video frame grabs, were found at depths> 2400 m, with a maximum of 3539 m while two of the others was collected at the much shallower depths of 963 and 919 m. The depth distribution of Tottonophyes enigmatica sp. nov. may have been affected by the actual depth of the water column, particularly at the deepest depths where specimens were usually found within 400 m of the sea bed. When the water depth was slightly shallower, as during Doc Ricketts Dive 105 and Tiburon Dive 897, the specimens were found very close to the seabed; being just 7 m above at the former, but the two shallowest specimens were collected at depths more than 600 m above the sea floor. Thus, it appears that T. enigmatica sp. nov. is largely restricted to deeper depths (> 2500 m), but may be upwelled to shallower depths within the complex canyon system in Monterey Bay. Little can be said about the relatively shallow depth of 550 m (water depth c. 2900 m) at which the strange eudoxid was collected until we can be sure that it actually belongs to T. enigmatica sp. nov. Molecular phylogenetics. The updated phylogenetic analyses for the Calycophorae presented here (Figure 13) builds directly on the early analysis of Dunn et al. (2005), and includes several additional calycophoran taxa. Tottonophyes enigmatica sp. nov. was included in the previously published analyses under the name Clausophyid sp. 1. The new analyses provide strong support for the monophyly of Calycophorae (Figure 13, branch 1) and also suggest that Prayomorpha is paraphyletic with respect to Diphyomorpha. Support for Diphyomorpha is moderate (Figure 13, branch 2). Tottonophyes enigmatica sp. nov. is placed as the sister taxon to the remaining Diphyomorpha, for which there is strong support (Figure 13, branch 3). Clausophyidae is recovered, but with very weak support. This indicates that Clausophyidae is either sister group to, or paraphyletic with respect to, the strongly supported clade indicated by branch 4 (Figure 13). This branch 4 clade consists of Sphaeronectes, Abylopsis, and all sampled taxa that have been assigned to Diphyidae. Etymology. We dedicate the family and hence the generic names to that great siphonophorologist Arthur Knyvett Totton who strove, in his Synopsis of the Siphonophora, to sort out many, but unfortunately not all, of the taxonomic problems that haunt this group. The specific name refers to the enigmatic combination of traits, some of which are unique to prayomorphs and others unique to diphyomorphs that until now had not been found in the same species. Discussion. The phylogenetic position of Tottonophyes enigmatica sp. nov. (Figure 13) and its unique combination of characters make it possible to postulate more specific hypotheses for the evolution of diphyomorph calycophorans than were possible in the past. These hypotheses can be described in terms of the character changes that occurred along four particularly interesting branches in the siphonophore phylogeny (Figure 13, branches 1–4). Changes along branch 1

Tottonophyes gen. nov. Monotypic genus for Tottonophyes enigmatica sp. nov., Published as part of Pugh, P. R., Dunn, C. W. & Haddock, S. H. D., 2018, Description of Tottonophyes enigmatica gen. nov., sp. nov. (Hydrozoa, Siphonophora, Calycophorae), with a reappraisal of the function and homology of nectophoral canals, pp. 452-472 in Zootaxa 4415 (3) on page 460, DOI: 10.11646/zootaxa.4415.3.3, http://zenodo.org/record/1242110

Agalma clausi (Bedot, 1888) Synonyms. Agalma sarsii (Fewkes, 1880) p. 628. Crystallodes Clausi (Bedot, 1888, post-scriptum) p. 20. Stephanopsis Clausi (Bedot, 1896) p. 406. Stephanomia sarsii (Schneider, 1898) p. 121–122. ? Agalma eschscholtzii (Haeckel, 1888) p. 226 and Plate XVIII, Fig. 8–17. ? Agalma haeckeli (Bigelow, 1911) p. 274–277. Systematics. Agalma clausi belongs to the Family Agalmatidae Brandt, 1834 that is nested within the historically recognized Physonectae, a suborder that groups all siphonophores sharing particular body plan i.e., possessing pneumatophore, nectosome, and siphosome. However, Physonectae were shown to be polyphyletic (Dunn et al. 2005) and following the most recent phylogeny (Munro et al. 2018) agalmatid siphonophores are placed within a newly erected, monophyletic clade Euphysonectae. Although this phylogenetic hypothesis is well supported, as the International Commission on Zoological Nomenclature (ICZN 1999) does not recognise clades we decided to maintain the name Physonectae throughout this text. Type specimens. There is no known type specimen, thus we designate the specimen from BWP Dive 1044-15 as a neotype. It will be deposited at the United States National Museum (Smithsonian Institution), Washington, DC under accession number USNM 1422474. It was collected close to the surface by a SCUBA diver on 14th July, 1983 at 3°56.9'N, 37°15.8'W. Diagnosis. Rigid-stemmed, monoecious agalmatid siphonophore of prismatic appearance, with nectophores arranged biserially and some bracts showing carmine pigmetation in living animal. Nectophores with short axial wings and size-dependent ridge pattern, incomplete laterals discernible only in younger zooids, but upper and lower lateral ridges present throughout development, but very weak in largest nectophores. No vertical lateral ridges. Thrust block small on mature nectophores; lateral ostial processes also small and packed with nematocysts. Typical Agalma course of lateral radial canals. Cormidia bearing gastrozooids (often flask-shaped), with tentacle, several gonophores of both sexes, bracts and palpons. Involucrum of tentillum voluminous, able to contain the cnidoband, the two terminal filaments and the central ampulla. Palpacle of palpon originates not at its base but, on the fully mature palpon, almost at its mid-length. Six types of bracts, of which one is able to discharge a coloured, probably fluorescent, fluid when stimulated., Published as part of Mańko, Maciej K. & Pugh, Philip R., 2018, Agalma clausi (Bedot, 1888) (Siphonophora: Physonectae) - complementary description with notes on species distribution and ecology, pp. 311-331 in Zootaxa 4441 (2) on pages 315-316, DOI: 10.11646/zootaxa.4441.2.7, http://zenodo.org/record/1301677, {"references":["Fewkes, J. W. (1880) Contributions to a knowledge of the tubular jellyfishes Bulletin of the Museum of Comparative Zoology at Harvard College, 7, 127 - 146, 3 pls.","Bedot, M. (1896) Les siphonophores de la Baie d'Amboine. Revue Suisse de Zoologie, 3, 367 - 414, 1 plate.","Schneider, K. C. (1898) Mittheilungen uber Siphonophoren. III. Systematische und andere Bemerkungen. Zoologischer Anzeiger, 21, 51 - 200.","Bigelow, H. B. (1911) The Siphonophorae. Memoirs of the Museum of Comparative Zoology at Harvard College, 38, 171 - 402. Brandt, J. F. (1834) Prodromus descriptionis animalium ab H. Mertensio in orbis terrarium circumnavigatione observatorum. Fascic. I. Polypos, Acalephas Discophoras et Siphonophoras, nec non Echinodermata continens. Recueil Actes des seances publiques de l'Acadademie imperiale des Science de St. Petersbourg, 1834, 201 - 275.","Munro, C., Siebert, S., Zapata, F., Howison, M., Damian-Serrano, A., Church, S. H., Goetz, F. E., Pugh, P. R., Haddock, S. H. D. & Dunn, C. W. (2018) Improved phylogenetic resolution within Siphonophora (Cnidaria) with implications for trait evolution. bioRxiv, 251116. https: // doi. org / 10.1101 / 251116","ICZN (1999) International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature, London, 306 pp."]}

A new species of calycophoran siphonophore, Tottonophyes enigmatica gen. nov, sp. nov., is described. It has a unique combination of traits, some shared with prayomorphs (including two rounded nectophores) and some with clausophyid diphyomorphs (the nectophores are dissimilar, with one slightly larger and slightly to the anterior of the other, and both possess a somatocyst). Molecular phylogenetic analyses indicate that the new species is the sister group to all other diphyomorphs. A new family, Tottonophyidae, is established for it. Its phylogenetic position and distinct morphology help clarify diphyomorph evolution. The function and homology of the nectophoral canals and somatocyst is also re-examined and further clarification is given to their nomenclature.

According to the latest phylogenetic and cytogenetic results, Larnax and Deprea should be merged in order to form a natural group. Consequently, we propose 32 combinations of Larnax species names under Deprea: D. abra-patriciae (S.Leiva & Barboza) S.Leiva & Deanna, comb. nov., D. altomayoensis (S.Leiva & Quip.) Barboza & Deanna, comb. nov., D. andersonii (N.W.Sawyer) Deanna & S.Leiva, comb. nov., D. bongaraensis (S.Leiva) Deanna & Barboza, comb. nov., D. chotanae (S.Leiva, Pereyra & Barboza) S.Leiva, comb. nov., D. darcyana (N.W.Sawyer) Barboza & S.Leiva, comb. nov., D. dilloniana (S.Leiva, Quip. & N.W.Sawyer) Barboza, comb. nov., D. grandiflora (N.W.Sawyer & S.Leiva) Deanna & Barboza, comb. nov., D. harlingiana (Hunz. & Barboza) S.Leiva & Deanna, comb nov., D. hawkesii (Hunz.) Deanna, comb. nov., D. kann-rasmussenii (S.Leiva & Quip.) S.Leiva & Barboza, comb. nov., D. longipedunculata (S.Leiva, E.Rodr. & J.Campos) Barboza, comb. nov., D. lutea (S.Leiva) Deanna, comb. nov., D. macasiana (Deanna, S.Leiva & Barboza) Barboza, comb. nov., D. maculatifolia (E.Rodr. & S.Leiva) S. Leiva, comb. nov., D. nieva (S.Leiva & N.W.Sawyer) Barboza & Deanna, comb. nov., D. parviflora (N.W.Sawyer & S.Leiva) S.Leiva, comb. nov., D. pedrazae (S.Leiva & Barboza) Deanna & S.Leiva, comb. nov., D. peruviana (Zahlbr.) S.Leiva & Barboza, comb. nov., D. pilosa (S.Leiva, E.Rodr. & J.Campos) Deanna, comb. nov., D. pomacochaensis (S.Leiva) Barboza, comb. nov., D. psilophyta (N.W.Sawyer) S.Leiva & Deanna, comb. nov., D. pumila (S.Leiva, Barboza & Deanna) S.Leiva, comb. nov., D. purpurea (S.Leiva) Barboza & S.Leiva, comb. nov., D. purpureocarpa (S.Leiva, Deanna & Barboza) Deanna, comb. nov., D. sachapapa (Hunz.) S.Leiva & Deanna, comb. nov., D. sagasteguii (S.Leiva, Quip. & N. W.Sawyer) Barboza, comb. nov., D. sawyeriana (S.Leiva, E.Rodr. & J.Campos) S.Leiva, comb. nov., D. schjellerupiae (S.Leiva & Quip.) Barboza & Deanna, comb. nov., D. steyermarkii (Hunz.) S.Leiva & Barboza, comb. nov., D. toledoana (Barboza & S.Leiva) Barboza, comb. nov., and D. vasquezii (S.Leiva, E.Rodr. & J.Campos) Deanna, comb. nov. Fil: Deanna, Rocío. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina Fil: Leiva González, Segundo. Universidad Privada Atenor Orrego. Museo de Historia Natural; Perú Fil: Barboza, Gloria Estela. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina

Kephyes hiulcus sp. nov. Kephyes sp. A. Grossmann & Lindsay 2013, Grossmann et al., 2015 Kephyes ovata. Pugh 2006 (Figs. 1, 5A) Kephyes ovata. Mapstone 2014 (Fig. 5D) Diagnosis. Anterior and posterior nectophores conical, ridgeless, laterally compressed. Nectosac extending to 90% height of nectophores, with looped radial canals. Lateral canals arise from upper canal, either together or separately. Pedicular canal with long descending diverticulum in both nectophores. Somatocyst thin, extending to nectophore apex. Hydroecium open over entire height of both nectophores, and without median mesogloeal protuberance from the hydroecial wall; hydroecial flaps ending slightly before ostial level, large hydroecial wings in posterior nectophore. Eudoxids cannot, at present, be differentiated from those of K. ovata. Material examined. Holotype: Polygastric stage comprised of attached anterior and posterior nectophores, and anterior part of siphosomal stem. Collected at 1189 m in Monterey Canyon by the ROV ��� Tiburon ��� (dive T0980- D 4, 10 May 2006, 36��36���11.909��� N, 122��22���31.998��� W). The holotype was deposited at the Showa Memorial Institute, National Science Museum, Tokyo (NSMT-Co1579). Additionally, twenty-one anterior nectophores and 25 posterior nectophores of Kephyes hiulcus sp. nov., ninety-seven anterior nectophores and 35 posterior nectophores of K. ovata, two complete Kephyes spp. free-living eudoxid stages, 302 bracts and 246 gonophores were analyzed, from Japanese waters and the Celebes and Mediterranean Seas (Table 1). Description of Holotype. Nectosome: Anterior nectophore (Figure 1): The type specimen was 12.6 mm in height and 3.6 mm wide in its preserved state. It was conical in shape, laterally flattened, without ridges; nectosac extending to 90% of nectophore height; no mouth plate; lateral radial canals arising from upper radial canal, with origins of right and left canals staggered, following a looped course, and inserting onto ostial ring canal in its lower lateral half; insertion point of pedicular canal at 40% nectophore height from the ostium; descending diverticulum from the pedicular canal remaining in contact with the hydroecium, extending at least to posterior end of hydroecial flaps; somatocyst remained in contact with hydroecium to 60% nectophore height before penetrating into the mesogloea, ending close to the apex of the nectophore; hydroecium open over the whole height of the nectophore, 20 to 45% of the nectophore width; hydroecial wings smooth and rounded, ending slightly anterior of the ostium, small projection of thickened mesogloea on external edge of hydroecial wings 1/3 nectophore height from the ostium. Posterior nectophore 12.9 mm in height and 4 mm wide in its preserved state; conical in shape, laterally flattened, without ridges; nectosac extended to 90% of the nectophore in height; no mouthplate; lateral radial canals arising together from upper radial canal, following a looped course, joining ostial ring canal in its lower lateral half; insertion point of pedicular canal 60% nectophore height from the ostium; descending branch of the pedicular canal remaining in contact with the hydroecium, extending at least to posterior end of hydroecial flaps; somatocyst remaining in contact with hydroecium for half its length before penetrating into the mesogloea, ending close to the apex of the nectophore; hydroecium open over the whole height of the nectophore; hydroecial wings large, ending slightly anterior of the ostium, with medial flaps on internal surface. Siphosome: Siphosomal cormidia composed of a bract and a tentacle-bearing gastrozooid (Figure 2A). No gonophores could be observed. The bracts could not be distinguished from those described by Pugh (2006) for Kephyes ovata, or from any of the material of both species we have examined. Gastrozooids pale tangerine in colour when alive. Cnidoband of tentacle tentilla long and slender (Figure 3). Although discharged nematocysts were not observed, the size and shape of the capsules suggested they were long thin microbasic mastigophores, measuring up to 112 ��m in length, and smaller, elongate homotrichous anisorhizas measuring up to 41 ��m in length. The terminal filament was covered in small, spherical nematocysts. Bracts measured up to 2.38 mm in length. Flattened apically, expanded laterally to grip the stem (Figure 2 B); neck-shield extensive, with small indentations on the posterior lateral corners; central phyllocyst spindle-shaped, extending to the apex of the headpiece; 2 longitudinal bracteal canals, up to 4/5 the neck-shield in length, sometimes curving sharply inwards in their most distal part, the right (when the neck-shield is downwards), being longer than the left. Gonophore not observed. Etymology. The specific name hiulcus is Latin meaning split or open, and refers to the wide-open or gaping hydroecium of the anterior nectophore of this species in comparison with the more-closed one of K. ovata. Distribution. Type locality: Monterey Canyon, Eastern Pacific Ocean (36��36���11.909��� N, 122��22���31.998��� W) Kephyes hiulcus sp. nov. samples have been collected in Monterey Canyon, in Japanese waters (Grossmann & Lindsay 2013, as ��� Kephyes sp. A���), in the Celebes Sea (Grossmann et al., 2015, as ��� Kephyes sp. A���), as well as in the Catalan Sea, north-western Mediterranean Sea (Table 1). The Kephyes hiulcus sp. nov. holotype was collected at 1189 m in Monterey Bay, but specimens from Japanese waters were found throughout mesopelagic depths. Nectophores were found to be roughly the same size, and were often collected concurrently with K. ovata. All of the MULTI-SPLASH net samples containing K. hiulcus sp. nov. also contained K. ovata, and 50% of the net samples from Suruga Bay contained both Kephyes species. There was no obvious segregation between the two species, or between polygastric and eudoxid stages, with depth, salinity or water temperature. The holotype was collected in waters with a temperature of 3.5˚C, salinity of 34.46 and a dissolved oxygen concentration of 0.56 mL.L -1, while the K. hiulcus sp. nov. specimens from the Catalan Sea (Mediterranean Sea) were collected in waters with temperatures above 13.0˚C, salinity 38.18���38.50 and dissolved oxygen concentrations between 4.14 and 5.87 ml.L -1. Kephyes ovata and K. hiulcus sp. nov. are the only members of the Clausophyidae for which the polygastric stage has been reported from the Mediterranean Sea. Remarks. First figured as a photograph (Fig. 5A: 1002) in Pugh���s (2006) reclassification of Clausophyes ovata (Keferstein & Ehlers, 1860) into the genus Kephyes, the present species K. hiulcus sp. nov. is similar in size, shape and general aspect to K. ovata. However, it can easily be differentiated by the extent of the hydroecium in the anterior nectophore of the polygastric stage. In K. ovata, this extends to 70% of the nectophore in height from the ostium (Pugh 2006), whereas in K. hiulcus sp. nov., the hydroecium is open over the entire height of the anterior nectophore. Additionally, the hydroecial wings do not extend to ostial level in K. hiulcus sp. nov., as they do in K. ovata, but end slightly anterior of the ostium, both in the anterior and posterior nectophores. In the posterior nectophores of K. hiulcus sp. nov., the anterior part of the somatocyst lies close to the apex of the nectophore, as in the anterior nectophore, while in K. ovata, the somatocyst does not reach the apex of the nectophore in the posterior nectophore. The origin of the lateral radial canals on the anterior nectophores of K. hiulcus sp. nov. showed a great deal of variability, arising either from the upper radial canal, or from the insertion point of the pedicular canal, right and left canals not always arising together. It is interesting to note that on all anterior K. ovata nectophores studied where the radial canals were visible, the lateral radial canals originated from the insertion point with the pedicular canal, as was the case in specimens from the north-eastern Pacific (Mapstone, 2009). Pugh (2006), when describing the variability of the origin of the lateral radial canal in K. ovata, used, by way of an example of the canals arising from the upper radial canal, the animal shown in his figure 5A (Pugh, 2006: 1002), which is, in fact, K. hiulcus sp. nov.. However, it is clear from Patriti���s (1969) specimens from the Mediterranean that in K. ovata the lateral radial canals may also arise separately, and from the upper radial canal. In Kephyes hiulcus sp. nov., neither the stem nor the posterior nectophore are attached level with the base of the somatocyst of the anterior nectophore, but rather some way down the hydroecial cavity. This can be clearly seen in the photograph of the living sampled holotype (Figure 1 A). Although no scar could be observed on the preserved material, it is supposed that the stem and posterior nectophore join at the start of the somatocyst of the latter, as in species of the genus Clausophyes, and that in K. hiulcus there is an extremely elongate ���somatocyst along the hydroecium��� somatocyst portion (Haddock et al. 2005) from the origin of the stem attachment point to the penetrating somatocyst of the anterior nectophore. The free-living eudoxid stages of the two Kephyes species cannot, at present, be differentiated. The eudoxid bracts of Kephyes spp. examined in the present study measured between 2.98 and 7.22 mm in length. The longitudinal canals were usually reduced, while the hydroecial canals extended to 15-20% of bract height from the posterior edge. On many bracts, the hydroecial canals ran in a smooth and slight curve towards the centre of the neckshield (Figure 2 D), resembling those drawn by Pugh for K. ovata (2006, Fig. 3A: p.1000). However, other bracts had hydroecial canals curving sharply inwards at their posterior ends, as observed on the small K. hiulcus sp. nov. siphosomal bracts (Figure 2 B), or again had one hydroecial canal bending inward at its posterior end, while the second had a bifurcated tip (Figure 2 C). The phyllocyst observed in free eudoxid stage bracts was more tapered in its anterior part than in the attached siphosomal bracts. It extended to or close to the apex of the head-piece (Figure 2). A well-developed hydroecial flap was present on the right hydroecial wing of all eudoxid stage gonophores studied. TABLE]. List of samples examineđ for the present stuđy. *: Kephyes hiulcus sp. nov. Holotype specimen; a.n.: anterior nectophore; b.: bract; g.: gonophore; e.: euđoxiđ; pol.: polygastric stage; p.n.: posterior nectophore. Species Zooid Depth Date Latitude Longitude Location Time Temperature (��C) Salinity Dissolved (No.) range (m) Oxygen (mL/L) Kephyes hiulcus sp. nov. pol.(1)* 1189 10-May-2006 36.603 -121.376 Monterey Canyon Day 3.52 34.46 0.6 p.n.(1) 0 0 750 21-Jun-2012 40.966 2.064 Catalan Sea Night 13.01023.0 7 38.18038.5 4.105. 9 p. n.(1) 0 0 750 21-Jun-2012 40.966 2.064 Catalan Sea Night 13.01023.0 7 38.18038.5 4.105. 9 p. n.(2) 90001000 19-Feb-2000 2.472 122.480 Celebes Sea Night 405 34.55034.6 1.801.9 a.n.(1) 7500800 15-Mar-2006 34.700 139.832 east of Oshima (Japan) Day 3.7604.0 0 34.32034.34 1.201.5 a.n.(1) 8000850 15-Mar-2006 34.700 139.832 east of Oshima (Japan) Day 3.5303.76 34.34034.37 1.201.4 a.n.(3) 8500900 22-Mar-2006 34.991 140.259 off Kamogawa (Japan) Day 3.3403.42 34.37034.38 1.101.2 a.n.(1) 4500500 23-Mar-2006 35.008 139.333 Sagami Bay Day 6.3407.18 34.25034.26 1.802.1 a.n.(6) 5000550 23-Mar-2006 35.008 139.333 Sagami Bay Day 5.7206.35 34.25034.25 1.701. 8 p. n.(3) 7500820 14-Mar-2006 35.008 139.333 Sagami Bay Night 3.6503.89 34.33034.36 1.301. 4 p. n.(1) 0 9-Mar-1980 25.200 124.600 Suruga Bay Night n/a n/a n/a a.n.(1) 397 27-Mar-1984 34.800 138.620 Suruga Bay Day 8.36 n /a n/a p.n.(2) 480 27-Nov-1986 34.850 138.637 Suruga Bay Day 6.88 n /a n/a p.n.(5) 494.1 9-Mar-1980 25.200 124.600 Suruga Bay Night n/a n/a n/a p.n.(1) 790 13-Nov-1992 34.858 138.633 Suruga Bay Night 3.94 34.36 n /a a.n.(1) 990 15-Mar-1985 40.487 142.658 Suruga Bay Day n/a n/a n/a a.n.(7) 1049 28-Mar-1985 34.795 138.617 Suruga Bay Day 3.16 n /a n/a a.n.(2) 1105 27-Nov-1986 34.850 138.637 Suruga Bay Day 3.93 n /a n/a p.n.(2) 1105 27-Nov-1986 34.867 138.638 Suruga Bay Night 3.93 n /a n/a a.n.(1) 4000450 8-Nov-1991 34.888 138.632 Suruga Bay Day 7.1606.73 34.257034.302 n/a p.n.(4) n/a 1980 n/a n/a Suruga Bay n/a n/a n/a n/a Kephyes ovata a.n.(1) 3500400 24-Mar-2006 34.700 139.832 east of Oshima (Japan) Day 7.0808.25 34.23034.28 2.302.6 a.n.(3) 3500400 25-Mar-2006 34.700 139.832 east of Oshima (Japan) Night 6.7008.18 34.22034.29 2.302.6 a.n.(1) 4000450 24-Mar-2006 34.700 139.832 east of Oshima (Japan) Night 6.47307.226 34.22034.26 2.202.4 a.n.(3) 4500500 24-Mar-2006 34.700 139.832 east of Oshima (Japan) Night 5.8206.48 34.20034.25 2.002.3 ������continued on the next page TABLE]. (Continueđ) Species Zooid Depth Date Latitude Longitude Location Time Temperature (��C) Salinity Dissolved (No.) range (m) Oxygen (mL/L) a.n.(1) 9000950 15-Mar-2006 34.700 139.832 east of Oshima (Japan) Day 3.2703.38 34.39034.41 0.901.2 ������continued on the next page TABLE]. (Continueđ) Species Zooid Depth Date Latitude Longitude Location Time Temperature (��C) Salinity Dissolved (No.) range (m) Oxygen (mL/L) a.n.(3) 480 27-Nov-1986 34.850 138.637 Suruga Bay Day 6.88 n /a n/a Kephyes spp. g.(1) 90001000 19-Feb-2000 2.472 122.480 Celebes Sea Night 4.0005.0 0 34.55034.60 1.801.9 ������continued on the next page TABLE]. (Continueđ) Species Zooid Depth Date Latitude Longitude Location Time Temperature (��C) Salinity Dissolved (No.) range (m) Oxygen (mL/L) b.(5) 5000550 24-Mar-2006 34.700 139.832 east of Oshima (Japan) Night 5.6005.85 34.19034.22 1.802.2 ������continued on the next page TABLE]. (Continueđ) Species Zooid Depth Date Latitude Longitude Location Time Temperature (��C) Salinity Dissolved (No.) range (m) Oxygen (mL/L) g.(4) 7000750 25-Mar-2006 34.991 140.259 off Kamogawa (Japan) Night 3.4804.10 34.29034.36 1.201.5 ������continued on the next page TABLE]. (Continueđ) Species Zooid Depth Date Latitude Longitude Location Time Temperature (��C) Salinity Dissolved (No.) range (m) Oxygen (mL/L) g.(14) 6000650 24-Mar-2006 35.008 139.333 Sagami Bay Night 4.6004.96 34.27034.28 1.501.6, Published as part of Grossmann, Mary Matilda & Lindsay, Dhugal John, 2017, A new species of clausophyid calycophoran siphonophore (Cnidaria: Hydrozoa), Kephyes hiulcus sp. nov., widely distributed throughout the world's oceans, pp. 43-54 in Zootaxa 4250 (1) on pages 44-53, DOI: 10.11646/zootaxa.4250.1.3, http://zenodo.org/record/439794, {"references":["Grossmann, M. M. & Lindsay, D. J. (2013) Diversity and distribution of the Siphonophora (Cnidaria) in Sagami Bay, Japan, and their association with tropical and subarctic water masses. Journal of Oceanography, 64 (4), 395 - 411. https: // doi. org / 10.1007 / s 10872 - 013 - 0181 - 9","Pugh, P. R. (2006) Reclassification of the clausophyid siphonophore Clausophyes ovata into the genus Kephyes gen. nov. Journal of the Marine Biological Association of the United Kingdom, 86, 997 - 1004.","Mapstone, G. M. (2014) Global diversity and review of Siphonophorae (Cnidaria: Hydrozoa). PloS ONE, 9 (2), 1 - 37. https: // doi. org / 10.1371 / journal. pone. 0087737","Mapstone, G. M. (2009) Siphonophora (Cnidaria: Hydrozoa) of Canadian Pacific waters. NRC Research Press, Ottawa, 302 pp.","Haddock, S. H. D., Dunn, C. W. & Pugh, P. R. (2005) A re-examination of siphonophore terminology and morphology, applied to the description of two new prayine species with remarkable bio-optical properties. Journal of the Marine Biological Association of the United Kingdom, 85, 695 - 707."]}

Genus Kephyes Pugh, 2006 Type species. Kephyes ovata (Keferstein & Ehlers, 1860). Diagnosis. Clausophyidae with two nectophores, lacking ridges and cusps, and with descending surface diverticula in both nectophores; cormidia with bracts, released as free-living eudoxids, each comprising a bract with two bracteal canals and a phyllocyst, a gastrozooid and tentacle, and a gonophore without distinctive ridges., Published as part of Grossmann, Mary Matilda & Lindsay, Dhugal John, 2017, A new species of clausophyid calycophoran siphonophore (Cnidaria: Hydrozoa), Kephyes hiulcus sp. nov., widely distributed throughout the world's oceans, pp. 43-54 in Zootaxa 4250 (1) on page 44, DOI: 10.11646/zootaxa.4250.1.3, http://zenodo.org/record/439794, {"references":["Pugh, P. R. (2006) Reclassification of the clausophyid siphonophore Clausophyes ovata into the genus Kephyes gen. nov. Journal of the Marine Biological Association of the United Kingdom, 86, 997 - 1004."]}

Family Clausophyidae Totton, 1965 Diagnosis. Calycophorae with dissimilar nectophores, typically two, both possessing a somatocyst and with anterior nectophore partially apposed to and partially linearly adjoined to the elongate posterior one; atypically only anterior one developed (genus Heteropyramis). Nectophore(s) either with longitudinal ridges or smooth; mouthplate absent in anterior nectophore (except in Clausophyes laetmata), present or absent in posterior nectophore; somatocyst in both nectophores typically simple, swollen, and reaching anterior end of nectophore. Cormidia with bracts, except in genera Clausophyes (eudoxids not developed) and Crystallophyes (produces a unique sexual stage termed the fuseudoxid); bract with phyllocyst variously shaped, with two fine basal branches (bracteal canals) extending down into neck-shield., Published as part of Grossmann, Mary Matilda & Lindsay, Dhugal John, 2017, A new species of clausophyid calycophoran siphonophore (Cnidaria: Hydrozoa), Kephyes hiulcus sp. nov., widely distributed throughout the world's oceans, pp. 43-54 in Zootaxa 4250 (1) on page 44, DOI: 10.11646/zootaxa.4250.1.3, http://zenodo.org/record/439794

Grossmann, Mary Matilda, Lindsay, Dhugal John (2017): A new species of clausophyid calycophoran siphonophore (Cnidaria: Hydrozoa), Kephyes hiulcus sp. nov., widely distributed throughout the world's oceans. Zootaxa 4250 (1): 43-54, DOI: https://doi.org/10.11646/zootaxa.4250.1.3

Erenna insidiator sp. nov. Diagnosis. Nectophores basic erennid ridge pattern. The upper laterals do not divide nor extend as far as the ostium. Two types of bract, both with pair of lateral cusps in proximal half; larger ones with rectangular distal ectodermal process on upper side, containing nematocysts. Very characteristic tentilla, consisting of a long pedicle and a long tube mostly filled with spongy tissue through which passes the endodermal canal. The tube tapered distally and there enclosing a photophore. Cnidoband itself forms the corrugated outer perimeter on a distinct process from the outside of the distal part of the cylinder. Material examined. Only two colonies of Erenna insidiator sp. nov. are known to have been collected and their station data are given here. Part of the specimen from Tiburon Dive 1040 is designated the holotype and will be donated to National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A., while the remainder, and the other specimen will be maintained, in the siphonophore collections at Brown University, Providence, R.I. Description. An in situ frame grab of the type specimen is shown in Figure 26, and a more detailed one of the recently collected specimen from Doc Ricketts Dive 779 is shown in Figure 27. Pneumatophore and Nectosome: In situ the nectosome of the type specimen occupied over half the total length with c. 29 mature nectophores and a number of nectophoral buds, together with the anterior pneumatophore. However, this region, having become denuded of its larger nectophores, does not appear to have been preserved. Nectophores: Twenty seven nectophores were found with the preserved type specimen that ranged in size from 12 to 30 mm in length and 6 to 29 mm in maximum width. The small nectophores (Figure 28 A) had yet to take on the flattened, in the upper/lower plane, appearance of the mature ones. The sides were folded up so that the upper lateral ridges ran parallel with each other and only a short distance apart. They defined the edges of a deep median furrow that was deepest proximally and gradually shallowed distally, as is indicated by the course of the upper radial canal in Figure 28 A. Proximally the combined upper and lower lateral ridges originated in the region where the nectophore was attached to the stem. The latter branched from the former at about half the height of the nectophore, and each was divided into two sections, proximal and distal, with the distal part appearing as an extension of the vertical lateral ridge (Figure 28 A). At this stage, the vertical lateral ridge did not join with the upper lateral ridge and, at all stages, neither the upper nor lower lateral ridges reached the ostium, nor did the former divide. The vertical and lower lateral ridges stood out from the surface of the nectophore. A small, elongate patch of ectodermal cells was present on each side of the nectophore just distal to the vertical lateral ridge. In the preserved state it was brown in colour but, as with other Erenna species, almost certainly it was black in life. The nectosac was deeply divided in the mid-line by the upper furrow (Figure 28 A), but had extensive lateral wings in the proximal half of the nectophore. The mantle canal was thick and U-shaped, and gave rise to the short pedicular canal that, in turn gave rise to all four radial canals. For the most part the lateral canals followed the contours of the nectosac, except distally where they continued to the lateral margins of the ostium. All the canals were a brownishblack in colour in the preserved state, but almost certainly were black in life. There was a small muscle-free zone just distal to the lateral radial canals, where they ascended from their origin from the pedicular canal. On the slightly larger nectophores (Figure 28 B, C) the upper lateral ridges still remained close together in the distal half of the nectophore, but the furrow between them was much shallower. The lower lateral ridge then appeared as a single entity and the vertical lateral ridges joined with the upper laterals. The muscle-free zone now consisted of a narrow band on either side of the proximal part of the lateral radial canals. The most interesting feature was the start of the development of the very characteristic thrust block. The central, proximal connection between the combined upper and lower lateral ridges, which formed the margins of the axial wings, consisted of two hemispherical mesogloeal thickenings that lay each side of a deep groove through which passed the ascending mantle canal. Distal to these, on the upper side of the nectophore, there were two further pairs of roughly hemispherical mesogloeal thickenings. The more distal pair was the larger and partially overhung the other pair. On the fully developed nectophores (Figure 29 A, B) the upper lateral ridges had now moved out toward the lateral margins. As before they did not quite reach to the ostium. The lower lateral ridges branched from the upper ones on the lateral margins of the axial wings, and these too did not quite reach ostial level. The vertical lateral ridges were distinct and there were small patches of, presumably, cells producing bioluminescence on the distal margins. Proximally the apex of the nectosac was almost flat and there was a clear muscle free zone, mainly on the lower side, but occasionally extending on to the upper one. The thrust block had expanded considerably and now formed two rounded lobes on the upper side of the nectophore. Below these and extending further proximally, was the pair of rounded thickenings that were deeply divided in the mid-line with the ascending mantle canal running through the furrow. The lateral ostial processes were well developed and even by the mature stage still contained nematocysts that appeared to be isorhizas (Figure 29 C). Siphosome: Because of the highly contracted nature of the preserved stem and the dense clusters of zooids, it was not possible to glean any useful information concerning the organisation of the individual cormidia. Bracts: Although very similar, the bracts could be divided into two types. The Type A (Figure 30 A) were slightly more abundant than the Type B ones (Figure 30 B). They were elongate, measuring up to c. 45 mm in length and 20 mm in width. They bore a pair of asymmetrically placed lateral cusps in their proximal half, with the cusp on the inner side usually being the more distal of the pair and more pronounced. There was a small circular patch of ectodermal cells on the upper side in the mid-line slightly distal of the mid-length. The inner side, at the proximal end, was distinctly thickened with mesogloea so that the bracteal canal ran over a sort of keel. The canal arose on the upper side of the bract and shortly after passing on to the lower side it usually thickened slightly. It continued distally and in contact with the lower wall until close to the distal end. There it thinned as it penetrated into the mesogloea and ran diagonally upwards to reach the upper surface toward the proximal end of a distinct raised patch of ectodermal cells, and then continued distally for a short distance immediately below the patch. Throughout its length the canal contained, in the preserved state, dark brown pigmentation that probably was black in life. This distal patch of cells (Figure 31) was roughly rectangular in shape with rounded corners. Its upper side consisted of a honeycomb of ectodermal cells interspersed with a few nematocysts. This probably was the result of abrasion as the more protected sides of the patch were packed with nematocysts. These nematocysts (Figure 31, insert) measured c. 100 x 28 ��m and were probably of the same type that were found on the lateral ostial processes of the nectophores. However, no evaginated ones were observed. The bracts of the second type (Figure 30 B) were shorter, measuring up to 25 mm in length and 17 mm in width. They also possessed a pair of lateral cusps in their proximal half, but in this case the cusps were almost symmetrically arranged. Again, there was a small central patch of ectodermal cells, and the course of the bracteal canal was the same as for the other type of bract. However, on the inner side of the bract, at the proximal end, the mesogloea was uniformly thickened and so there was no pronounced keel. In the very youngest bracts there was a conical protuberance on one side of the bract, proximally, but this disappeared as the bracts enlarged. The distal ectodermal thickening on the upper side of the bract was basically the same as for the other type of bract, although in general it was less extensive. Gastrozooid and tentacle: The large gastrozooids (Figure 32) were largely black in colour and, in both life and after preservation, had an asymmetrical shape. The small, ring-like basigaster was attached directly to the stem and was separated from the main stomach region by an annulation. The stomach was usually laterally expanded and the distal proboscis region arose at an oblique angle on one side. The distal region was paler than the others and numerous endodermal, longitudinal "hepatic stripes" could be seen. The tentacle arose from one side of the basigaster, and was annulated and bore strong musculature. Tentillum: The tentillum (Figure 33) was of a unique morphology, although typical for an Erenna species. It consisted of an obviously highly extensile, but in its preserved state a highly contracted pedicle, as exemplified by the corrugated nature of the gastrovascular cavity. The remainder of the tentillum was a cylindrical tube, but flattened on its inner side. It was U-shaped in its preserved state and measured c. 55 mm in length, but in life was quite straight (see Figure 27). Its ectoderm consisted of irregularly shaped small cells, below which was an amorphous layer of mesogloea, and then a spongy internal layer of pentagonal or hexagonal-shaped cells surrounding the narrow gastrovascular cavity (Figure 33 B). This spongy layer decreased in size distal to the cnidoband itself, forming a narrower, short terminal process that terminated in a spherical photophore (Figure 34 D). There the gastrovascular cavity gave rise to a diverticular canal that ran proximally, above the main gastrovascular cavity to the middle of the cnidoband (Figure 33 C, D). Whether it then gave rise to two branches that ran distally and proximally along the base of the cnidoband could not be discerned. Young tentilla, at various stages in development are shown in Figure 34 A. The outer process that forms the cnidoband is distinct at a very early stage in development. No details with regard to the ocellus and its potential luminescent properties are presently available. The cnidoband itself, as with other Erenna species (see Pugh, 2001) was a discrete structure that measured c. 12.5 mm in length in its preserved state (Figure 33 B, C). The nematocysts were concentrated around the outer lateral margins of the cnidoband in a much folded sequence giving the outer edge a scalloped appearance. The large nematocysts, presumably microbasic mastigophores (Figure 36 B), which measured c. 142 x 23 ��m, were present in relatively large numbers compared with the other Erenna species. The smaller ones (Figure 34 C), which were quite variable in size, measuring c. 39��� 42 x 16���20 ��m, appeared to be all of the ovoid, pear-shaped type. No discharged ones of either type were found. Whether the small cylindrical type of nematocyst was absent or so rare that it was not noted was unclear. Palpon: Each palpon (Figure 35) consisted of a short pedicle, a slightly broader part, separated from the more distal parts by a constriction, to which the palpacle was attached. The latter was often lost, but when present, was very short and possessed no nematocysts. The main part of the palpon consisted of a large, flimsy bag-like structure, together with a distinct, narrow, whitish proboscis region. The main body was covered in an array of long irregular patches of large ectodermal cells interspersed with or overlaying more homogeneous smaller cells. The largest palpons measured up to c. 20 mm in length and 10 mm in diameter. Groups of palpons arose from a common pedicle and these were usually closely associated with the gonodendra. However, the highly contracted nature of the preserved stem did not allow for the exact arrangement to be elucidated. From 30 to 50 nematocysts were found around the distal mouth opening that measured c. 92 x 28 ��m (Figure 35 inset). Although of similar size to those found on the nectophores and bracts they did not appear to be of the same type. Possibly they were mastigophores, but this could not be confirmed as no discharged ones were found. Gonodendron: The type specimen was male and gonophores, in various stages of development, were found attached to several gonodendra (Figure 36 A). There appeared to be more than one of these per cormidium, but due to the highly contracted state of the preserved specimen, this could not be verified with certainty. Each gonodendron appeared to be associated with a cluster of palpons, but again the highly contracted nature of the preserved siphosome did not allow an accurate assessment of their disposition. The younger gonophores were clustered closely together, but as they matured they became more separated, being borne on rigid, brittle, almost twig-like, stalks. Each gonophore, up to c. 3.5 mm in diameter, was a spherical, bag-like structure with a somewhat amorphous but spherical, central manubrium containing the sexual products that occupied only part of the subumbrella space. Typically four straight radial canals could be discerned, but they were not arranged symmetrically (see Figure 36 B). Their arrangement suggested that there should have been five canals, spaced approximately 72�� apart, but with one canal missing. Occasionally (see Figure 36 B) this fifth canal was present, while on rare occasions there were only three canals that reached the ostial ring canal. The ostium was completely closed by a circular membrane comprised of an outer ring of large, roughly circular cells surrounding smaller cells. Distribution. Known from only two specimens, one collected from a depth of 3098 m at 34��17.23'N, 124��03.10'W, west of Point Conception, California, where the water depth was c. 3450 m, and the other west of the Davidson Seamount, California at a depth of 2800 m, where the water depth was c. 3770 m. Etymology. The specific name insidiator is derived from the Latin insidia, and means an ambusher as, although not proven, its tentillum probably, in some way, acts as a lure., Published as part of Pugh, P. R. & Haddock, S. H. D., 2016, A description of two new species of the genus Erenna (Siphonophora: Physonectae: Erennidae), with notes on recently collected specimens of other Erenna species, pp. 401-446 in Zootaxa 4189 (3) on pages 424-432, DOI: 10.11646/zootaxa.4189.3.1, http://zenodo.org/record/166117, {"references":["Pugh, P. R. (2001) A review of the genus Erenna Bedot, 1904 (Siphonophora, Physonectae). Bulletin of the Natural History Museum, London (Zoology Series), 67, 169 - 182."]}