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The swimming larvae of many marine animals identify a location on the seafloor to settle and undergo metamorphosis based on the presence of specific surface-bound bacteria. While bacteria-stimulated metamorphosis underpins processes such as the fouling of ship hulls, animal development in aquaculture, and the recruitment of new animals to coral reef ecosystems, little is known about the mechanisms governing this microbe-animal interaction. Here we review what is known and what we hope to learn about how bacteria and the factors they produce stimulate animal metamorphosis. With a few emerging model systems, including the tubeworm Hydroides elegans, corals, and the hydrozoan Hydractinia, we have begun to identify bacterial cues that stimulate animal metamorphosis and test hypotheses addressing their mechanisms of action. By understanding the mechanisms by which bacteria promote animal metamorphosis, we begin to illustrate how, and explore why, the developmental decision of metamorphosis relies on cues from environmental bacteria.

Larvae of many marine invertebrates bear an anteriorly positioned apical sensory organ (ASO) presumed to be the receptor for settlement- and metamorphosis-inducing environmental cues, based on its structure, position and observed larval behavior. Larvae of the polychaeteHydroides elegansare induced to settle by bacterial biofilms, which they explore with their ASO and surrounding anteroventral surfaces. A micro-laser was utilized to destroy the ASO and other anterior ciliary structures in competent larvae ofH. elegans. After ablation, larvae were challenged with bacterial biofilmed or clean surfaces and percent metamorphosis was determined. Ablated larvae were also assessed for cellular damage by applying fluorescently tagged FMRF-amide antibodies and observing the larvae by laser-scanning confocal microscopy. While the laser pulses caused extensive damage to the ASO and surrounding cells, they did not inhibit metamorphosis. We conclude that the ASO is not a required receptor site for cues that induce metamorphosis.Summary StatementLarvae of the polychaeteHydroides elegansretain the capacity to sense biofilm cues and metamorphose despite removal of their apical sensory organs, the supposed sensors for settlement cues.

Made available in DSpace on 2019-10-06T16:40:54Z (GMT). No. of bitstreams: 0 Previous issue date: 2019-09-01 Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) Human occupation of coastal areas promotes the establishment of non-native species but information on bioinvasions is usually biased toward the Northern Hemisphere. We assessed non-native species' importance in sessile communities at six marinas along the most urbanized area of the Southwestern Atlantic coastline. We found 67 species, of which 19 are exotic. The most frequent species was the exotic polychaete Branchiomma luctuosum, while the most abundant was the exotic bryozoan Schizoporella errata that monopolized the substrata in three marinas. Along with S. errata, the exotic polychaete Hydroides elegans and ascidian Styela plicata dominated space in the three remaining marinas, while native species were in general rare. We show that communities associated with artificial substrata along this Brazilian urbanized area are dominated by exotic species and that using abundance data along with species identity can improve our understanding of the importance of exotic species for the dynamics of biological communities. Centro de Ciências Naturais e Humanas Universidade Federal do ABC (UFABC), Rua Arcturus, 03 - Jardim Antares Departamento de Zoologia Instituto de Biociências Universidade de São Paulo, R. Matão, Trav. 14 Museu Nacional Universidade Federal do Rio de Janeiro (UFRJ), Quinta da Boa Vista, s/n – São Cristóvão Departamento de Biologia Marinha Universidade Federal Fluminense Universidade Federal do Rio de Janeiro Instituto de Biologia Departamento de Zoologia, Av. Carlos Chagas Filho, 373 Departamento de Biologia Animal Instituto de Biologia Universidade Estadual de Campinas (UNICAMP), Caixa Postal 6109 Laboratório de Estudos de Bryozoa – LAEBry Departamento de Zoologia Centro de Biociências Universidade Federal de Pernambuco (UFPE), Av. Prof. Moraes Rego 1235 - Cidade Universitária Laboratório de Evolução e Diversidade Aquática – LEDA Departamento de Ciências Biológicas Universidade Estadual Paulista (UNESP), FCL/Assis, Av. Dom Antônio, 2100 Departamento de Zoologia Universidade Federal do Paraná Laboratório de Evolução e Diversidade Aquática – LEDA Departamento de Ciências Biológicas Universidade Estadual Paulista (UNESP), FCL/Assis, Av. Dom Antônio, 2100 CAPES: 001 FAPESP: 2015/24408-4 FAPESP: 2016/17647-5 FAPESP: 2016/50389-0 CNPq: 308768/2018-3 CNPq: 309995/2017-5

Fouling polychaetes in Tanjung Priok Port, Jakarta and their status were studied in order to update the list of marine alien species in Indonesia. Polychaetes were collected using six PVC panels that were submerged at a depth of 0.5 m over three months in Jakarta Bay. Polychaetes were identified to the lowest taxon possible based on characteristics of external morphology. The geographical distribution of identified polychaetes was then determined based on taxonomic literature in order to justify whether they were native or alien species for Indonesian waters. Twelve fouling polychaetes belonging to nine families are reported here. Of these, nine species (i.e., Chrysopetalum debilis, Eunice hirschi, Ceratonereis mirabilis, Leonnates decipiens, Polyophthalmus pictus, Eulalia (Eumida) sanguinea, Lepidonotus tenuisetosus, Hydroides elegans, and Polydora ciliata) were native species, while three other taxa (i.e., Namanereis sp., Hypsicomus sp. 1, and Hypsicomus sp. 2) were undetermined.

A 14-month study was conducted of the polychaetous annelids present on attached wooden blocks at nine stations in Los Angeles Harbor in 2013-2014. The results were compared to a similar study conducted at the same stations in 1950-1951. The primary objective in both studies was to determine the location and occurrence of marine borers in the harbor. Since fouling organisms, including polychaetes, attached to the wooden blocks, it also provided an opportunity to study the polychaetes that settled on the blocks. The number of polychaete species in these two studies increased from 22 to 71. The serpulid Hydroides elegans was a dominate species in both studies but the pollution indicator Capitella capitata, common in the earlier study, was rare in the recent study. There was a seasonal occurrence in both the number of species and specimens with highs in the warmer months and lows from December through March in both studies. Many environmental changes have occurred in the Los Angeles Harbor complex over the past 63 years. New harbor piers constructed that extend into the Outer Harbor, channels have been dredged deeper and pollution abatement programs initiated. The water quality has been improved especially in the Inner Harbor as a result of these changes where the dissolved oxygen in the water was low or absent in 1950-1951 but higher (over 6.0 mg/L) in 2013-2014. This study is unusual since long-term, seasonal comparisons of marine invertebrate populations are uncommon.

The swimming larvae of many marine animals identify a location on the sea floor to undergo metamorphosis based on the presence of specific bacteria. Although this microbe–animal interaction is critical for the life cycles of diverse marine animals, what types of biochemical cues from bacteria that induce metamorphosis has been a mystery. Metamorphosis of larvae of the tubeworm Hydroides elegans is induced by arrays of phage tail-like contractile injection systems, which are released by the bacterium Pseudoalteromonas luteoviolacea. Here we identify the novel effector protein Mif1. By cryo-electron tomography imaging and functional assays, we observe Mif1 as cargo inside the tube lumen of the contractile injection system and show that the mif1 gene is required for inducing metamorphosis. Purified Mif1 is sufficient for triggering metamorphosis when electroporated into tubeworm larvae. Our results indicate that the delivery of protein effectors by contractile injection systems may orchestrate microbe–animal interactions in diverse contexts.

Ocean uptake of anthropogenic CO2 causes ocean acidification (OA), which not only decreases the calcification rate, but also impairs the formation of calcareous shells or tubes in marine invertebrates such as the dominant biofouling tubeworm species, Hydroides elegans. This study examined the ability of tubeworms to resume normal tube calcification when returned to ambient pH 8.1 from a projected near-future OA level of pH 7.8. Tubeworms produced structurally impaired and mechanically weaker calcareous tubes at pH 7.8 compared to at pH 8.1, but were able to recover when the pH was restored to ambient levels. This suggests that tubeworms can physiologically recover from the impacts of OA on tube calcification, composition, density, hardness and stiffness when returned to optimal conditions. These results help understanding of the progression of biofouling communities dominated by tubeworms in future oceans with low pH induced by OA.

The serpulid Hydroides elegans is recognized as an invasive species in many harbour areas of the world, growing mainly on man-made structures such as pier pilings, where native diversity is low. In addition, because they build large aggregations of calcareous tubes, this increases the maintenance and fuel costs of vessels. While H. elegans is well known for the North Atlantic, Europe, Caribbean and Pacific Ocean, detailed information of the species for the South Atlantic is missing. In the last decade, owners of boats moored at the Clube Naval Charitas (CNC), in the Guanabara Bay, southeastern Brazil noticed an increase in the occurrence of an encrusting organism that we identified as H. elegans. In this study we characterize H. elegans specimens from CNC and Itaguai Harbor, both in southeastern Brazil, and discuss its distribution in the world. In addition, we deployed PVC panels at two different depths to quantify how H. elegans affects community assembling over two months. We observed that H. e...

Hydroides elegans (Haswell, 1883) Figs 4E, 5 Eupomatus elegans Haswell, 1883: 633, pl. 12, fig. 1 (type locality: Port Jackson [= Sydney], Australia). Eupomatus pectinatus Philippi, 1844: 195, pl. 6, fig. R (type locality: Mediterranean). Hydroides abbreviata Kr��yer in M��rch, 1863: 377, pl. 11, figs 6���7 (type locality: Saint Croix Island, Lesser Antilles, Caribbean Sea). Hydroides pacificus Hartman, 1969: 759���760, figs 1���5 (type locality: Velero IV, sta. 1454���42, from ship hull; central and southern California). Hydroides norvegica (non Gunnerus 1768) ��� Edmonson & Ingram 1939: 268���271 (Kaneohe Bay and Pearl Harbor, Oahu, Hawaii; pier fouling). ��� Edmonson 1944: 3���16 (Pearl Harbor, Oahu, Hawaii; fouling experiments; 0���4 m). ��� Berkeley & Berkeley 1941: 56 (Newport Bay, southern California; from piling). ��� Hartman 1952: 63���64, figs 1���2 (Corpus Christi Bay, Texas; bottom of a boat; some specimens with double funnel). ��� Renaud 1956: 35 (Miami, Florida; bottom of R/V Physalia). ��� Hartman 1961: 44 (Los Angeles harbor, southern California; fouling on hulls of ships). ��� Rioja 1961: 311 (Ant��n Lizardo Beach, Veracruz, eastern Mexico; on ascidians and shells). ��� Lakshmana Rao 1969: 5, pl. 2, figs a���g (Visakhapatnam and Madras, now Chennai, India; harbours). Serpula vermicularis (non Linnaeus 1767) ��� Lakshmana Rao 1969: 2���3, pl. 1, figs a���g (Visakhapatnam and Madras, now Chennai, India; harbours). Hydroides elegans ��� Zibrowius 1971: 721���725, figs 56���64 (Mediterranean, Western Africa, South Africa, Mozambique, Java, Australia, Hawaii, California, Florida; extensive revision; intertidal to 6 m; on hull of boats). ��� Long 1974: 28 (Pearl Harbor, Oahu, Hawaii; 9 m; with little coverage on fouling plates). ��� Bailey-Brock 1976: 77���78 (Oahu Island and Hawaii Island; reef flats, on chlorophyte Dictyosphaeria cavernosa (Forssk��l) B��rgesen, epifauna of mobile substrata (mollusks and crustaceans), boat harbors and lagoons, brackish waters and reef slope). ��� Imajima 1976b: 237���238, fig. 3a���n (Tokyo Bay, Kanagawa Prefecture, Shizuoka Prefecture, Mie Prefecture, Wakayama Prefecture, Osaka Bay, Hiroshima Bay, Nagasaki Harbour, Amakusa Bay and Kagoshima Bay, Kochi Prefecture, Koniya Island, Japan; on the oysters Pinctada fucata (Gould, 1850), Crassostrea gigas (Thunberg, 1793) and Pteria penguin (R��ding, 1798)); 1979: 169 (around Shionomisaki Cape and Kushimoto Harbour, southern Japan; 19���75 m; on shell, gravel and rope). ��� Due��as 1981: 100���101, fig. 31A���G (Cartagena Bay, Colombia; on plastic pond related to shrimp culture). ��� Imajima 1982: 46 (Arumizu Bay, Palau Islands). ��� Bailey-Brock 1987: 420��� 421 (Hawaii). ��� Zibrowius 1992: 91 (discussion about its origin). ��� Nishi 1995a: 101���102 (southern Japan; on chelae and carapace of the crab Charybdis riversandersoni Alcock, 1899). ��� Ishaq & Mustaquim 1996: 170, fig. 5A���H (Karachi, Pakistan; intertidal, on rocks, buoy and boat hull). ��� Nishi 1996: 306���308 (Okinawa Island, Southwest Japan; buried in the coral Montastrea sp. and attached to dead coral, Pocillopora damicornis (Linnaeus, 1758)). ��� Perkins 1998: 95 (checklist of shallow-water polychaetes from Florida). ��� Due��as 1999: 14 (Cartagena Bay, Colombia). ��� Bastida-Zavala & Salazar-Vallejo 2000b: 846���848, fig. 2e���f (eastern Mexico: San Juan de Ul��a, Veracruz; Champot��n, Campeche; Celest��n, Yucat��n; and Contoy Island, Quintana Roo; 0.3���3.5 m; on calcareous rocks, seagrass and macroalgae, wood dock pilings, and fort wall cover with vermetids, oysters and ascidians). ��� D��az-D��az & Li��ero-Arana 2001: 12 (Cariaco Gulf, Venezuela; PVC dock pilings). ��� Bastida-Zavala & ten Hove 2002: 164���166, figs 35A���J, 36 (Florida, Texas, Puerto Rico, Atlantic Panama, Aruba, Cura��ao, and eastern Mexico: Veracruz, Campeche and Quintana Roo; intertidal to 7 m; 31���37���; on coral debris, fouling of wood pier and boats). ��� Bastida-Zavala & ten Hove 2003: 86���87, fig. 11A���S (California and Hawaii; intertidal to 1 m; among algae and bryozoans, fouling of marina piers and ship and submarine hulls). ��� Rodr��guez-Valencia 2004: 520 (Petacalco Bay, Guerrero, southern Mexican Pacific; 3���21 m). ��� ��inar 2006: 226���227, fig. 3D���E (as NIS from eastern Levantine coast of Turkey). ��� Bastida-Zavala 2008: 25���26, fig. 6H (California and Baja California Sur, Mexican Pacific; intertidal to 1 m,on PVC plates and hull of boats).��� Carlton & Eldredge 2009: 62 (Hawaii; invasion history). ��� D��az-Casta��eda &Valenzuela-Solano 2009: 513 (west coast of Baja California Peninsula:Salsipuedes Bay, Baja California; vicinity of tuna farm sea-cages). ��� Tovar-Hern��ndez et al. 2009: 331, figs 3l, 8a-c (as fouling species in Mazatl��n, Sinaloa, Mexican Pacific). ��� Ben-Eliahu & ten Hove 2011: 19��� 25, figs 5D���E, 6���7 (Turkey, Cyprus, Israel, Egypt, Suez Canal and Red Sea; 0.2���10 m, on algae (Cystoseira myrica C. Agardh now Polycladia myrica (S.G. Gmelin) Draima, Ballesteros, F.Rousseau & T. Thibaut, Digenea, Laurensia and Sargassum), sponges, gastropods (Murex forskoehli R��ding, 1798), bivalves (Brachidontes pharaonis, Chama gryphoides Linnaeus, 1758, Chicoreus erythraeus, now Murex erythraeus P. Fischer, 1870, Crenatula picta, Fulvia fragilis (Forssk��l in Niebuhr, 1775), Fusinus verrucosus (Gmelin, 1791), Malvufundus normalis (Lamarck, 1819), now Malleus anatinus (Gmelin, 1791), M. regulus (Forssk��l in Niebuhr, 1775), Pinctada radiata, Spondylus spinosus Schreibers, 1793 and Pectinidae), bryozoans, barnacles, crabs, tunicates, under rocks, artificial substrates such as canal walls, a tin can submerged in mud, rubber fenders and iron frames). ��� Tovar-Hern��ndez et al. 2012: 16���17 (Gulf of California: Guaymas, Sonora and Topolobampo, Sinaloa); 2014: 388, 390 (Gulf of California: Topolobampo, Sinaloa; Guaymas, Sonora; La Paz, Baja California Sur). ��� Schwan et al. 2015: 3���6, fig. 2A���I (southeastern Brazil; intertidal to 1 m; fouling of harbors, marinas and PVC plates). ��� Sun et al. 2015: 23���29, fig. 6a���b (New South Wales, Northern Territory, Queensland, South and Western Australia; intertidal to 20 m; on scallop and mussel clumps on sandy bottom, fine mud, under rocks, unvegetated sediment, Zostera, Caulerpa filiformis, brown algae, on barnacles, cement and wooden pilings, floating pontoon, fouling plates, inside air-conditioning cooling pipe, and ship hulls). ��� Bastida-Zavala et al. 2016: 419���420, figs 4, 11F���G (Mexican Pacific: Baja California Sur and Oaxaca; in marinas and harbors, fouling; intertidal to 1 m). Hydroides pacificus ��� D��az-Casta��eda 2000: 327 (west coast of Baja California Peninsula: Todos Santos Bay, Baja California; 10 m; terracota plates). Material examined 384 specimens: IR (123) Aug. 2005, BB (1) Aug. 2004, TB (32) Jun. and Jul. 2012, HB (1) Sep. 2003, LB (87) Sep. 2003, MI (5) Jul. 2013, SD (87) Sep. 2000 and Jul. 2013, HI (48) Aug. 2006. Additional material 18 specimens: LACMNH s.n., colony (southern California, 33��21'03" N, 118��19'55" W, off Avalon, Santa Catalina Island, Velero III, sta. 1377, grey sand, 110 m, 3 Aug. 1941, as Serpula sp.); LACMNH s.n., 1 specimen (southern California, 33��44'02" N, 118��32'03" W, 10 km from Pt Vicente lighthouse, Velero IV, sta. 2475, peel grab, mud, specimen attached to hexactinellid sponge, 740 m, 28 Oct. 1953); LACMNH s.n., 1 specimen (southern California, 33��47'59" N, 118��33'59" W, 11.5 km from Palos Verdes Point, Velero IV, sta. 2792, Hayward grab, mud, 600 m, 22 May 1954); LACMNH s.n., 1 specimen (southern California, 33��14' N, 118��18'04" W, 6.9 km from East End Light, Santa Catalina Island, Velero IV, sta. 2850, Campbell grab, 1200 m, 23 Jun. 1954); MBL-SD s.n., 5 specimens (southern California, approx. 32��43' N, 117��13' W, San Diego, CSA P���161, 1 month plate, 7 Oct. to 7 Nov. 1974, as H. pacificus). Diagnosis This species is gregarious and can form small colonies. Tube white, with or without two longitudinal ridges, with or without peristomes; but not alveoli. Opercular peduncle smooth, white. Opercular funnel with 15���31 radii (21���35 in eastern Pacific specimens) with blunt tips (Fig. 4E); verticil with 11���18 spines, straight, with pointed tips (Fig. 4E); all spines with 0���4 internal spinules and 2���3 pairs of lateral spinules (2���5 in eastern Pacific specimens), without external spinules and/or wings (Fig. 4E). Special collar chaetae with two pointed-elongate teeth and a proximal rasp, distal blade with notch and many denticles. Taxonomic remarks Hydroides elegans now exhibits a world-wide distribution in tropical and subtropical ports, marinas and eutrophic lagoons (ten Hove 1974). Due to its rapid colonization and population growth, it is considered an invasive species (Zibrowius 1973, 1992, 1994; Tovar-Hern��ndez et al. 2009, 2014). The main means of dispersal of this species is through fouling on boats and ships and/or as larvae in ballast water (Nelson-Smith 1967; Tovar-Hern��ndez et al. 2009). Although H. elegans was described from Australia (Haswell 1883), and Zibrowius (1992) and several authors speculate that it is native there (Ben-Eliahu & ten Hove 2011; Sun et al. 2015), this is not clear evidence of an Australian origin. Bastida-Zavala et al. (2016: 420) draw attention to the fact that both the Hawaiian (Long 1974; Bailey-Brock 1976) and Australian (Sun et al. 2015) records of this species were from both natural and man-made substrates, suggesting a broader Indo-West Pacific origin. However, there are also records of this species from both natural and artificial substrates in the Gulf of Mexico and Caribbean Sea (e.g., Hartman 1952; Bastida-Zavala & Salazar-Vallejo 2000b; Bastida-Zavala & ten Hove 2002). In addition, 20 years before H. elegans was described, Kr��yer (in M��rch 1863) described H. abbreviata, a senior synonym of the former species, from Saint Croix, a Caribbean Island, suggesting a possible Caribbean origin as well. Finally, the oldest description of a senior synonym of H. elegans is H. pectinatus, described by Philippi in 1844, from the Mediterranean. One of these old names could take precedence over the name H. elegans (Principle of Priority, ICZN 1999, art. 23); however, nomenclatural stability will not promote the re-introduction of these oldest available names, and therefore the conditions for reversal of precedence are met (ICZN 1999, art. 23.9). To resolve the origin of this species, a molecular analysis of the different world-wide populations will be necessary. In this work, Hydroides elegans is considered a cryptogenic species. Ecology Intertidal to sublittoral (10 m). Some records off California (additional material from LACMNH) have questionable depth data (110���1200 m). In subtropical and tropical marine and brackish waters; salinities of 31���37��� (Bastida-Zavala & ten Hove 2002); on reef flats, on chlorophytes, epifauna of mobile substrata (mollusks and crustaceans), boat harbors, lagoons and reef slope (Bailey-Brock 1976); on the oysters Pinctada fucata, Crassostrea gigas and Pteria (Magnavicula) penguin (Imajima 1976b); on the crab Charybdis riversandersoni, dead coral (Pocillopora damicornis) and buried in the coral Montastrea sp. (Nishi 1995 a, 1996); also on rocks, seagrass, macroalgae (Cystoseira myrica, Digenea, Laurencia and Sargassum), sponges, gastropods (Murex forskoehli), bivalves (Brachidontes pharaonis, Chama gryphoides, Chicoreus erythraeus, Crenatula picta, Fulvia fragilis, Fusinus verrucosus, Malvufundus normalis, M. regulus, Pinctada radiata, Spondylus spinosus and Pectinidae), bryozoans, barnacles, crabs, tunicates, coral debris, under rocks, and on artificial substrates such as boat and ship hulls, cement and wood dock pilings, buoys, PVC or terracotta plates, canal walls, tin cans, rubber fenders and iron frames (Hartman 1952; Long 1974; D��az-Casta��eda 2000; Bastida-Zavala & Salazar-Vallejo 2000b; Bastida-Zavala & ten Hove 2002, 2003; Ben-Eliahu & ten Hove 2011). Distribution Worldwide in tropical to temperate waters. Western Atlantic: Caribbean Sea, Bermuda, Gulf of Mexico, Brazil; eastern Pacific: Hawaii, southern California, Mexican Pacific, Panama; other regions: Mediterranean, Suez Canal, western and South Africa, Mozambique, Pakistan, Java, Hong Kong, Australia, southern and central Japan, Palau Islands (M��rch 1863; Fauvel 1932; Day 1967; Zibrowius 1971; Imajima 1976b, 1979, 1982; Ishaq& Mustaquim 1996; Bastida-Zavala & ten Hove2002, 2003; ��inar 2006; Bastida-Zavala 2008; Ben-Eliahu & ten Hove 2011; Sun et al. 2012, 2015). In this work, Hydroides elegans was found abundantly and frequently on fouling plates from the Indian River and Tampa Bay, Florida; Long Beach and San Diego Bay, California; and Oahu, Hawaii; and occasionally from Biscayne Bay, Florida, and Humboldt Bay and Mission Bay, California (Fig. 5). Only one specimen was found on a fouling plate from Humboldt Bay, California, in 2003, suggesting that the species may now be present in northern California. However, additional sampling is necessary to confirm that H. elegans is currently established in northern California. Hydroides elegans was observed at Eel Pond, Cape Cod, Massachusetts, on one occasion in the summer and fall of 2011, surviving the winter 2011���2012, and found again in the spring of 2012 occurring at remarkable abundances. However, after the second winter, no live specimens were found (Fofonoff et al. 2003; Jim Carlton, pers. comm., 2011���2013) so it is not clear if the species is still established there. If so, this record extends the northward range of the species on the east coast, from the Indian River, Florida, to Cape Cod, Massachusetts (1750 km). The remarkable abundance observed by Carlton demonstrates the colonization potential of H. elegans with warming temperatures, such as those predicted under the climate change scenario. On the east coast, Eel Pond has a latitude of 41��33' N, while on the west coast, Humbolt Bay, northern California, is located at 40��45' N; both localities represent the northernmost records of this species on mainland North America, suggesting that continued monitoring is necessary to evaluate the species spread., Published as part of Bastida-Zavala, J. Rolando, McCANN, Linda D., Keppel, Erica & Ruiz, Gregory M., 2017, The fouling serpulids (Polychaeta: Serpulidae) from United States coastal waters: an overview, pp. 1-76 in European Journal of Taxonomy 344 on pages 30-34, DOI: 10.5852/ejt.2017.344, http://zenodo.org/record/3834679, {"references":["Haswell W. A. 1883. On some new Australian tubicolous Annelids. Proceedings of the Linnean Society of New South Wales 7: 633 - 638.","Philippi A. 1844. Einige Bemerkungen uber die Gattung Serpula, nebst Aufzahlung der von mir im Mittelmeer mit dem Thier beobachteten Arten. Archiv fur Naturgeschichte 10 (1): 186 - 198. https: // doi. org / 10.5962 / bhl. part. 29558","Morch O. A. L. 1863. Revisio critica Serpulidarum. Et bidrag til rorormenes naturhistorie. Naturhistorisk Tidsskrift, Series 3, 1: 347 - 470.","Hartman O. 1969. Atlas of the Sedentariate Polychaetous Annelids from California. Allan Hancock Foundation, Los Angeles.","Gunnerus J. E. 1768. Om nogle Norske Coraller. Det Kongelige Norske Videnskabens Selskab Skrifter 4: 38 - 73.","Edmonson C. H. & Ingram W. M. 1939. Fouling organisms in Hawaii. Occasional Papers of the Bernice P. Bishop Museum 14 (14): 251 - 300. http: // hbs. bishopmuseum. org / pubs-online / pdf / op 14 - 14. pdf","Edmonson C. H. 1944. Incidence of fouling in Pearl Harbor. Occasional Papers of the Bernice P. Bishop Museum 18 (1): 1 - 34. http: // hbs. bishopmuseum. org / pubs-online / pdf / op 18 - 1. pdf","Berkeley E. & Berkeley C. 1941. On a collection of Polychaeta from Southern California. Bulletin of the Southern California Academy of Sciences 40 (1): 16 - 60.","Hartman O. 1952. Fouling serpulid worms, new to the Gulf of Mexico. Texas Journal of Sciences 1: 63 - 64.","Renaud J. C. 1956. A report on some polychaetous annelids from the Miami-Bimini area. American Museum Novitates 1812: 1 - 40. http: // digitallibrary. amnh. org / handle / 2246 / 4653","Hartman O. 1961. Polychaetous annelids from California. Allan Hancock Pacific Expeditions 25: 1 - 226.","Rioja E. 1961. Estudios Anelidologicos XXIV: Adiciones a la fauna de anelidos poliquetos de las costas orientales de Mexico. Anales del Instituto de Biologia 31: 289 - 316.","Lakshmana Rao M. V. 1969. Fouling serpulids from some Indian harbours. Journal of the Timber Development Association of India 15 (2): 1 - 20.","Linnaeus C. 1767. Systema Naturae. 12 th ed., 1, Part 2. L. Salvius, Holmiae [Stockholm], 533 - 1327. Linero-Arana I. & Diaz-Diaz O. 2012. Presencia del poliqueto exotico Ficopomatus uschakovi (Polychaeta: Serpulidae) en Venezuela: descripcion y comentarios sobre su distribucion. Interciencia 37: 234 - 237.","Zibrowius H. 1971. Les especes Mediterraneennes du genre Hydroides (Polychaeta Serpulidae): remarques sur le pretendu polymorphisme de Hydroides uncinata. Tethys 2: 691 - 746.","Long E. R. 1974. Marine fouling studies off Oahu, Hawaii. The Veliger 17 (1): 23 - 36.","Bailey-Brock J. H. 1976. Habitats of tubicolous polychaetes from the Hawaiian Islands and Johnston Atoll. Pacific Science 30 (1): 69 - 81. http: // scholarspace. manoa. hawaii. edu / bitstream / handle / 10125 / 1532 / v 30 n 1 - 69 - 81. pdf","Imajima M. 1976 b. Serpulidae (Annelida, Polychaeta) from Japan. I. The genus Hydroides. Bulletin of the National Science Museum, Series A (Zoology) 2 (4): 229 - 247.","Duenas P. D. 1981. Inventario preliminar de los poliquetos (Annelida) de aguas someras de la Bahia de Cartagena y areas adyacentes. Boletin del Museo del Mar (10): 82 - 138.","Imajima M. 1982. Serpulinae (polychaetous annelids) from the Palau and Yap Islands, Micronesia. Proceedings of the Japanese Society of Systematic Zoology 23: 37 - 55.","Bailey-Brock J. H. 1987. Phylum Annelida. In: Devaney D. M. & Eldredge L. G. (eds) Reef and Shore Fauna of Hawaii. Section 3: Sipuncula through Annelida: 213 - 453. Special Publication of the Bernice P. Bishop Museum, 64.","Zibrowius H. 1992. Ongoing modification of the Mediterranean marine fauna and flora by the establishment of exotic species. Mesogee 51: 83 - 107.","Nishi E. 1995 a. Occurrence of the serpulid polychaetes, Hydroides elegans (Haswell) and H. fusicola Morch on the carapace of the crustaceans. Nankiseibutu: The Nanki Biological Society 37 (2): 101 - 102.","Ishaq S. & Mustaquim J. 1996. Polychaetous annelids (order Sabellida) from the Karachi coast, Pakistan. Pakistan Journal of Marine Sciences 5 (2): 161 - 197. http: // aquaticcommons. org / 17776 / 1 / PJMS 5.2 _ 161. pdf","Nishi E. 1996. Serpulid polychaetes associated with living and dead corals at Okinawa Island, Southwest Japan. Publications of the Seto Marine Biological Laboratory 37 (3 / 6): 305 - 318. https: // repository. kulib. kyoto-u. ac. jp / dspace / bitstream / 2433 / 176262 / 1 / fia 0373 - 6 _ 305. pdf","Perkins T. H. 1998. Checklist of shallow-water marine polychaetous Annelida of Florida. In: Camp D. K., Lyons W. G. & Perkins T. H. (eds) Checklist of Selected Shallow-water Marine Invertebrates of Florida: 79 - 122. Florida Marine Research Institute, Technical Report 3.","Duenas P. D. 1999. Algunos poliquetos (Annelida) del Caribe colombiano. Milenio 1 (2): 9 - 18.","Bastida-Zavala J. R. & Salazar-Vallejo S. I. 2000 b. Serpulidos (Polychaeta: Serpulidae) del Caribe noroccidental: Hydroides y Serpula. Revista de Biologia Tropical 48: 841 - 858.","Diaz-Diaz O. & Linero-Arana I. 2001. Poliquetos asociados a substratos artificiales sumergidos en la costa nororiental de Venezuela. II: Serpulidae y Spirorbidae. Boletin del Instituto de Oceanografia de Venezuela 40 (1 - 2): 9 - 20.","Bastida-Zavala J. R. & ten Hove H. A. 2002. Revision of Hydroides Gunnerus, 1768 (Polychaeta: Serpulidae) from the Western Atlantic Region. Beaufortia 52: 103 - 178.","Bastida-Zavala J. R. & ten Hove H. A. 2003. Revision of Hydroides Gunnerus, 1768 (Polychaeta: Serpulidae) from the Eastern Pacific Region and Hawaii. Beaufortia 53: 67 - 110.","Rodriguez-Valencia J. A. 2004. Respuesta de los poliquetos bentonicos a la variabilidad ambiental y condiciones El Nino en Bahia Petacalco (Guerrero, Mexico). Ciencias Marinas 30 (4): 515 - 526. http: // libros. duhnnae. com / 2017 / may / 14944091826 - Respuesta-de-los-poliquetos-bentonicos-a-lavariabilidad-ambiental-y-condiciones-el-nino-en-bahia. php","Cinar M. E. 2006. Serpulid species (Polychaeta: Serpulidae) from the Levantine coast of Turkey (eastern Mediterranean), with special emphasis on alien species. Aquatic Invasions 1 (4): 223 - 240. https: // doi. org / 10.3391 / ai. 2006.1.4.6","Bastida-Zavala J. R. 2008. Serpulids (Annelida: Polychaeta) from the eastern Pacific, including a brief mention of Hawaiian serpulids. Zootaxa 1722: 1 - 61.","Carlton J. T. & Eldredge L. G. 2009. Marine Bioinvasions of Hawai'i. The Introduced and Cryptogenic Marine and Estuarine Animals and Plants of the Hawaiian Archipelago. Bishop Museum Bulletins in Cultural and Environmental Studies 4. Bishop Museum Press, Honolulu.","Diaz-Castaneda V. & Valenzuela-Solano S. 2009. Polychaete fauna in the vicinity of bluefin tuna seacages in Ensenada, Baja California, Mexico. Zoosymposia 2: 505 - 526.","Tovar-Hernandez M. A., Mendez N. & Villalobos-Guerrero T. F. 2009. Fouling polychaete worms from the southern Gulf of California: Sabellidae and Serpulidae. Systematics and Biodiversity 7 (3): 319 - 336. https: // doi. org / 10.1017 / S 1477200009990041","Ben-Eliahu M. N. & ten Hove H. A. 2011. Serpulidae (Annelida: Polychaeta) from the Suez Canal- From a Lessepsian migration perspective. Zootaxa 2848: 1 - 147. http: // www. mapress. com / zootaxa / 2011 / f / zt 02848 p 147. pdf","Tovar-Hernandez M. A., Villalobos-Guerrero T. F., Yanez-Rivera B., Aguilar-Camacho J. M. & Ramirez- Santana I. D. 2012. Guia de Invertebrados acuaticos exoticos en Sinaloa. Geomare, A. C., USFWS, INE- SEMARNAT, Mazatlan, Sinaloa, Mexico.","Schwan I. D. S., Brasil A. C. D. S., Neves D. & Dias G. M. 2015. The invasive worm Hydroides elegans (Polychaeta - Serpulidae) in southeastern Brazil and its potential to dominate hard substrata. Marine Biology Research 12 (1): 96 - 103. https: // doi. org / 10.1080 / 17451000.2015.1080370","Sun Y., Wong E., ten Hove H. A., Hutchings P. A., Williamson J. E. & Kupriyanova E. K. 2015. Revision of the genus Hydroides (Annelida: Serpulidae) from Australia. Zootaxa 4009 (1): 1 - 99. https: // doi. org / 10.11646 / zootaxa. 4009.1.1","Bastida-Zavala J. R., Rodriguez Buelna A. S., de Leon-Gonzalez J. A., Camacho-Cruz K. A. & Carmona I. 2016. New records of sabellids and serpulids (Polychaeta: Sabellidae, Serpulidae) from the Tropical Eastern Pacific. Zootaxa 4184 (3): 401 - 457. https: // doi. org / 10.11646 / zootaxa. 4184.3.1","Diaz-Castaneda V. 2000. The early establishment and development of a polychaete community settled on artificial substrata at Todos Santos Bay, Baja California, Mexico. Bulletin of Marine Science 67 (1): 321 - 335.","Hove H. A. ten. 1974. Notes on Hydroides elegans (Haswell, 1883) and Mercierella enigmatica Fauvel, 1923, alien serpulid polychaetes introduced into the Netherlands. Bulletin Zoologisch Museum, Universiteit van Amsterdam 4 (5): 45 - 51.","Zibrowius H. 1973. Remarques sur trois especes de Serpulidae acclimatees en Mediterranee: Hydroides dianthus (Verrill, 1873), Hydroides dirampha Morch, 1863, et Hydroides elegans (Haswell, 1883).","Zibrowius H. 1994. Introduced invertebrates: examples of success and nuisance in the European Atlantic and in the Mediterranean. In: Boudouresque C. F., Briand F. & Nolan C. (eds) Introduced Species in European Coastal Waters: 44 - 49. Ecosystems Research Report 8, European Commission.","Tovar-Hernandez M. A., Yanez-Rivera B., Villalobos-Guerrero T. F., Aguilar-Camacho J. M. & Ramirez- Santana I. D. 2014. Invertebrados marinos exoticos en el Golfo de California. In: Low Pfeng A. M, Quijon P. A. & Peters Recagno E. M. (eds) Especies invasoras acuaticas: Casos de Estudio en Ecosistemas de Mexico: 15 - 32. Secretaria de Medio Ambiente y Recursos Naturales, Instituto Nacional de Ecologia y Cambio Climatico, Mexico DF, Mexico, and University of Prince Edward Island, Charlotteville, Canada.","Nelson-Smith A. 1967. Catalogue of Main Marine Fouling Organisms (found on Ships coming into European Waters). Vol. 3. Serpulids. Organization for Economic Co-operation and Development, Paris.","ICZN (International Commission on Zoological Nomenclature). 1999. International Code of Zoological Nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London.","Fossa S. A. & Nielsen A. J. 1996. Einzellige Organismen, Schwamme, marine Wurmer und Weichtiere im Korallenriff und fur das Korallenriff-Aquarium. In: Korallenriff-Aquarium 5. Birgit Schmettkamp Verlag, Bornheim, Germany.","Fauvel P. 1932. Annelida Polychaeta of the Indian Museum, Calcutta. Memoirs of the Indian Museum 12: 1 - 262. http: // faunaofindia. nic. in / PDFVolumes / memoirs / 012 / 01 / 0001 - 0262. pdf","Day J. H. 1967. A Monograph on the Polychaeta of South Africa. Part 2: Sedentaria. British Museum (Natural History), London.","Imajima M. 1979. Serpulidae (Annelida, Polychaeta) collected around Cape Shionomisaki, Kii Peninsula. Memoirs of the National Science Museum 12: 159 - 183.","Sun Y., ten Hove H. A. & Qiu J. W. 2012. Serpulid polychaetes from Hong Kong. Zootaxa 3424: 1 - 42.","Fofonoff P. W., Ruiz G. M., Steves B. & Carlton J. T. 2003. National Exotic Marine and Estuarine Species Information System [online]. Available from http: // invasions. si. edu / nemesis / [accessed 22 Jul. 2016]."]}

Larvae of many sessile marine invertebrates settle in response to surface microbial communities (biofilms), but the effects of soluble compounds from biofilms in affecting larval behavior prior to settlement, attachment, and metamorphosis have been little studied. This question was addressed by videotaping the behavior of competent larvae of the serpulid, Hydroides elegans, above settlement-inducing biofilms. Adult worms were collected in Pearl Harbor, Hawaii, USA in November 2012 and spawned almost immediately. Six-day old larvae were placed in five replicated treatments in small cups: (1) with a natural biofilm; (2) with a natural biofilm on an 8-µm screen, 1 mm above the bottom of a clean cup; (3) with a natural biofilm beneath a clean screen; (4) in a clean cup; and (5) in a clean cup with a clean screen. Using the videotapes, larval swimming speeds and trajectories were quantified within 5 min of the larvae being placed in a treatment. Only larvae that touched a biofilm, i.e., in treatments (1) and (2), slowed their swimming speed and increased the amount of time spent crawling rather than swimming. This shows that under these conditions, any soluble cues emanating from a biofilm do not affect settlement behavior. Furthermore, after 24 h close to 100 % of larva in the two accessible biofilm treatments had metamorphosed and

Hydroides elegans (Haswell, 1883) (Figures 4, 11 F���G) Eupomatus elegans Haswell, 1883: 633, pl. 12, Fig. 1. Type locality: Port Jackson, Australia. Hydroides norvegica (not Gunnerus, 1768).��� Berkeley & Berkeley 1941: 56 (Newport Bay, Southern California, from ���piling���); Hartman 1961: 44 (Los Angeles harbor, Southern California, fouling on hulls of ships); Lakshmana Rao 1969: 5, pl. 2, Figs a���g (Visakhapatnam and Madras, India; harbours); Salazar-Vallejo & Londo��o-Mesa 2004: 54 (Tropical Eastern Pacific, checklist). Hydroides pacificus Hartman, 1969: 759 ���760, Figs 1���5 (type locality: Velero IV, sta. 1454-42, from hull of ship; central and Southern California); D��az-Casta��eda 2000: 327 (Todos Santos Bay, Baja California, terracotta plates, 10 m). Serpula verimicularis (not Linnaeus, 1767).��� Lakshmana Rao 1969: 2 ���3, pl. 1, Figs a���g (Visakhapatnam and Madras, India; harbours; auctore misspelling). Hydroides elegans.��� Zibrowius 1971: 721���725, Figs 56���64 (Oahu, Hawaii, Newport Bay and Los Angeles Harbor, California); Long 1974: 28 (Pearl Harbor, Oahu, Hawaii, 9 m, fouling, with little coverage on test panels); Bailey-Brock 1976: 77���78 (Oahu Island and Hawaii Island, reef flats, live substrata [chlorophyte Dictyosphaeria cavernosa], epifauna of mobile substrata [mollusks and crustaceans], boat harbors, lagoons, brackish waters and reef slope); Due��as 1981: 100��� 101, fig. 31A���G (Cartagena Bay, Colombia; on plastic ponds for shrimp culture); Bailey-Brock 1987: 420���421 (Hawaii); Zibrowius 1992: 91 (discussion about its origin); Carpizo-Ituarte & Hadfield 1998: 15, Fig. 2 (Pearl Harbor, Hawaii, stimulation of metamorphosis); Bastida-Zavala & ten Hove 2003: 86���87, Figs 11 A���S (California and Hawaii; 0���1 m); Rodr��guez-Valencia 2004: 520 (Petacalco Bay, Guerrero); Okolodkov et al. 2007: 40 (cryptogenic in the Mexican Pacific); Bastida-Zavala 2008: 25���26, fig. 6H (California and Baja California Sur; 0���1 m; deeper records questionable); Bastida- Zavala 2009: 535, Figs 2 E, 5F (identification key for Tropical America); D��az-Casta��eda & Valenzuela-Solano 2009: 513 (Salsipuedes Bay, Baja California, near tuna sea-cages); ten Hove & Kupriyanova 2009: 53 (worldwide serpulid checklist); Tovar-Hern��ndez et al. 2009b: 331, Figs 3 l, 8a-c (fouling in Mazatl��n, Sinaloa); Tovar-Hern��ndez et al. 2012: 16���17 (Guaymas, Sonora, and Topolobampo, Sinaloa); Bastida-Zavala et al. 2014: 326, Figs 19.1e ���h (exotic in the Mexican Pacific); Tovar-Hern��ndez et al. 2014: 390 (Marina Palmira, Topolobampo, Sinaloa; API and Marina Fonatur, Guaymas, Sonora; and API and Marina Palmira, La Paz, Baja California Sur); Villalobos-Guerrero et al. 2014: 107 (Sinaloa, checklist); Sun et al. 2015: 23���29, fig. 6a���b (Capital Territory, New South Wales, Northern Territory, Queensland, South Australia and Western Australia; intertidal to 20 m; in natural habitat as lagoons, also in fouling communities of fish farms, harbors and ship���s hulls). Material examined. 3,545 specimens. Baja California Sur: UANL 7875, 548 spec. (Marina Santa Rosal��a, sta. 3: 27��20���24.5���N, 112��15���56.1���W, April 19, 2010, ARB & JAL); UANL 7876 (same, sta. 4: 27��20���23.9���N, 112��15���55.9���W, April 19, 2010, ARB & JAL); UANL 7877, 1,271 spec. (same, Sta. 1, 27��20���25.2���N, 112��15���56.1���W, March 31, 2011, coll. JAL & ARB); UANL 7878, 1,702 spec. (Puerto Escondido, sta. 1: 25��48���51.8���N, 111��18���41.2���W, sta. 2: 25��48���53.1���N, 111��18���40.5���W, April 2, 2011, coll. ARB & JAL); UANL 7879 (Marina Palmira, La Paz, 24��11���05.3���N, 110��18���12.8���W, April 3, 2011, coll. ARB & JAL). Oaxaca: UMAR-Poly 765, 4 spec. (Salina Cruz, angler pier, main dock, sta. 4, 1 m, May 26, 2011, coll. SGM et al.); UMAR-Poly 766, 10 spec. (same, hull of the shrimp boat ���Golfo P��rsico���, 1 m, May 26, 2011, coll. EVP): UMAR-POLY 767, 8 spec. (���sample 85���, Oaxaca, 0���6 m, September 15, 2004). Habitat. Intertidal to subtidal (20 m, Sun et al. 2015); deeper records, 110��� 1,200 m from California (LACM- AHF 1377, 2475, 2792, 2850), were regarded to be questionable by Bastida-Zavala (2008: 26). Most records of Hydroides elegans were as fouling on artificial substrates: hulls of ships, terracotta and PVC panels and harbor structures. In the Hawaiian Islands it was also found on reef flats, reef slope and on the native alga Dictyosphaeria cavernosa (Bailey-Brock 1976). In the material studied the specimens were associated only with anthropogenic substrates in marinas and ports from La Paz, Puerto Escondido and Santa Rosal��a, Baja California Sur, and from Salina Cruz, Oaxaca. Fouling species. Distribution. Worldwide in temperate and tropical regions. Mediterranean, North Sea, Gulf of Mexico to Brazil, South Africa, Persian Gulf, India, Australia, Micronesia; California (USA) to Oaxaca (M��xico), Hawaii (Zibrowius 1971; Bastida-Zavala & ten Hove 2002; 2003; Sun et al. 2015). Remarks. The confusion between Hydroides elegans and H. norvegica Gunnerus, 1768 was unraveled by Zibrowius (1971), mainly on the basis of the collar chaetae: in H. norvegica with 2���3 teeth, while they have many small teeth in H. elegans. Hydroides norvegica has a distribution limited to the boreal Atlantic and subtidal waters in the Mediterranean, while H. elegans has a worldwide distribution in temperate and tropical waters, mainly in harbors and marinas (Zibrowius 1971; ten Hove 1974; Zibrowius 1992; Kupriyanova & Jirkov 1997; Bastida- Zavala & ten Hove 2002; 2003; Moen 2006; Bastida-Zavala 2008; Ben-Eliahu & ten Hove 2011). Long (1974: 28) recorded both H. elegans and H. norvegica from Oahu, Hawaii; however, after the revision of Zibrowius (1971) it is uncertain if the specimens identified by Long as H. norvegica belong to H. elegans or a different taxon. Zibrowius (1992) doubted if Hydroides elegans originated from Australia; however, Ben-Eliahu & ten Hove (2011) propose that its origin probably is Australia and Sun et al. (2015) suggest on biogeographical and ecological reasons that H. elegans is likely to be a native to Australia. Many historic and recent records of H. elegans come from ports and marinas, on anthropogenic substrates; however, in the Hawaiian Islands the species was found in both natural sites and in boat harbors (Long 1974; Bailey-Brock 1976); also, H. elegans was found in Australia both in natural sites (lagoons), as well as fouling of fish farms, harbors and ship���s hulls (Sun et al. 2015). Hydroides elegans has been present in the Eastern Pacific for at least 86 years; its first record was Newport Bay, Southern California (Berkeley & Berkeley 1941, as Hydroides norvegica); Bastida-Zavala & ten Hove (2003) recorded specimens collected in 1929 from the fouling of a submarine (U.S. Narwhal, N-1) in the harbor of San Francisco. In the Tropical Eastern Pacific the species was recorded first by Bastida-Zavala (2008) with a sample of more than a thousand specimens encrusting (1991) a PVC plate in La Paz Bay, Baja California Sur; later, Tovar- Hern��ndez et al. (2009b) recorded the species in the fouling of Mazatl��n port, Sinaloa. Hydroides elegans was selected as a model species for research in biofouling and testing marine coatings for a number of reasons: its high abundance and the fact that their calcareous tubes can create problems for vessels. Furthermore H. elegans is easy to reproduce in laboratory (Carpizo-Ituarte & Hadfield 1998; Nedved & Hadfield 2008), making it possible to experiment with different means to avoid their settlement, e.g. antifouling paints (Johnson & Gonzalez 2004; Johnson et al. 2006); and has been observed that the larvae settle in response to both natural biofilms, formed by marine bacteria, or to artificially induced surfaces by cations and other chemicals (Nedved & Hadfield 2008)., Published as part of Bastida-Zavala, J. Rolando, Buelna, Alondra Sof��a Rodr��guez, Le��n-Gonz��lez, Jes��s Angel De, Camacho-Cruz, Karla Andrea & Carmona, Isabel, 2016, New records of sabellids and serpulids (Polychaeta: Sabellidae, Serpulidae) from the Tropical Eastern Pacific, pp. 401-457 in Zootaxa 4184 (3) on pages 419-420, DOI: 10.11646/zootaxa.4184.3.1, http://zenodo.org/record/255062, {"references":["Haswell, W. A. (1883) On some new Australian tubicolous Annelids. Proceedings of the Linnean Society of New South Wales, 7, 633 - 638.","Gunnerus, J. (1768) Om nogle norske coraller. Skrifter det Kongliger norske Videnskabsselskabet Trondhjem, 4, 38 - 73.","Berkeley, E. & Berkeley, C. (1941) On a collection of Polychaeta from Southern California. Bulletin of the Southern California Academy of Sciences, 40 (1), 16 - 60.","Hartman, O. (1961) Polychaetous annelids from California. Allan Hancock Pacific Expeditions, 25, 1 - 226.","Lakshmana Rao, M. V. (1969) Fouling serpulids from some Indian harbours. Journal of Timber Developers Association of India, 15 (2), 1 - 20.","Salazar-Vallejo, S. I. & Londono-Mesa, M. H. (2004) Lista de especies y bibliografia de poliquetos (Polychaeta) del Pacifico oriental tropical. Anales del Instituto de Biologia, Mexico, 75 (1), 9 - 97.","Hartman, O. (1969) Atlas of the Sedentariate Polychaetous Annelids from California. Allan Hancock Foundation, Los Angeles, 812 pp.","Diaz-Castaneda, V. (2000) The early establishment and development of a polychaete community settled on artificial substrata at Todos Santos Bay, Baja California, Mexico. Bulletin of Marine Science, 67 (1), 321 - 335.","Zibrowius, H. (1971) Les especes Mediterranee du genre Hydroides (Polychaeta Serpulidae). Remarques sur le pretendu polymorphisme de Hydroides uncinata. Tethys, 2, 691 - 746.","Long, E. R. (1974) Marine fouling studies off Oahu, Hawaii. The Veliger, 17 (1), 23 - 36.","Bailey-Brock, J. H. (1976) Habitats of tubicolous polychaetes from the Hawaiian Islands and Johnston Atoll. Pacific Science, 30 (1), 69 - 81.","Duenas, P. D. (1981) Inventario preliminar de los poliquetos (Annelida) de aguas someras de la bahia de Cartagena y areas adyacentes. Boletin del Museo del Mar, 10, 82 - 138.","Bailey-Brock, J. H. (1987) Phylum Annelida. In: Devaney, D. M. & Eldredge, L. G. (Eds.), Reef and shore fauna of Hawaii. Section 3: Sipuncula through Annelida. Special Publication of the Bernice P. Bishop Museum, 64, pp. 213 - 453.","Zibrowius, H. (1992) Ongoing modification of the Mediterranean marine fauna and flora by the establishment of exotic species. Mesogee, 51, 83 - 107.","Carpizo-Ituarte, E. & Hadfield, M. G. (1998) Stimulation of metamorphosis in the polychaete Hydroides elegans Haswell (Serpulidae). The Biological Bulletin, 194 (1), 14 - 24. http: // dx. doi. org / 10.2307 / 1542509","Bastida-Zavala, J. R. & Hove, H. A. ten (2003) Revision of Hydroides Gunnerus, 1768 (Polychaeta: Serpulidae) from the Eastern Pacific Region and Hawaii. Beaufortia, Bulletin of Zoological Museum, 53 (4), 67 - 110.","Rodriguez-Valencia, J. A. (2004) Respuesta de los poliquetos bentonicos a la variabilidad ambiental y condiciones El Nino en Bahia Petacalco (Guerrero, Mexico). Ciencias Marinas, 30 (4), 515 - 526.","Okolodkov, Y. B., Bastida-Zavala, R., Ibanez, A. L., Chapman, J. W., Suarez-Morales, E., Pedroche, F. & Gutierrez-Mendieta, F. J. (2007) Especies acuaticas no indigenas en Mexico. Ciencia y Mar, 11 (32), 29 - 67.","Bastida-Zavala, J. R. (2009) Serpulidae Rafinesque, 1815. In: Leon-Gonzalez, J. A. de, Bastida-Zavala, J. R., Carrera-Parra, L. F., Garcia-Garza, M. E., Pena-Rivera, A., Salazar-Vallejo, S. I. & Solis-Weiss, V. (Eds.), Poliquetos (Annelida: Polychaeta) de Mexico y America Tropical. Vol. 4 3. Universidad Autonoma de Nuevo Leon, Monterrey, pp. 521 - 554.","Diaz-Castaneda, V. & Valenzuela-Solano, S. (2009) Polychaete fauna in the vicinity of bluefin tuna sea-cages in Ensenada, Baja California, Mexico. Zoosymposia, 2, 505 - 526.","Hove, H. A. ten & Kupriyanova, E. K. (2009) Taxonomy of Serpulidae (Annelida, Polychaeta): the state of affairs. Zootaxa, 2036, 1 - 126.","Tovar-Hernandez, M. A., Mendez, N. & Villalobos-Guerrero, T. F. (2009 b) Fouling polychaete worms from the Southern Gulf of California: Sabellidae and Serpulidae. Systematics and Biodiversity, 7 (3), 319 - 336. http: // dx. doi. org / 10.1017 / s 1477200009990041","Tovar-Hernandez, M. A., Yanez-Rivera, B., Villalobos-Guerrero, T. F., Aguilar-Camacho, J. M. & Ramirez-Santana, I. D. (2014) Invertebrados marinos exoticos en el Golfo de California. In: Low Pfeng, A. M, Quijon, P. A. & Peters Recagno, E. M. (Eds.). Especies invasoras acuaticas: Casos de estudio en ecosistemas de Mexico. Secretaria de Medio Ambiente y Recursos Naturales, Instituto Nacional de Ecologia y Cambio Climatico, University of Prince Edwards Island, Prince Edward Island, pp. 15 - 32.","Bastida-Zavala, J. R. & Hove, H. A. ten (2002) Revision of Hydroides Gunnerus, 1768 (Polychaeta: Serpulidae) from the Western Atlantic Region. Beaufortia, 52, 103 - 178.","Hove, H. A. ten (1974) Notes on Hydroides elegans (Haswell, 1883) and Mercierella enigmatica Fauvel, 1923, alien serpulid polychaetes introduced into the Netherlands. Bulletin Zoologisch Museum Universiteit van Amsterdam, 4 (5), 45 - 51.","Kupriyanova, E. K. & Jirkov, I. A. (1997) Serpulidae (Annelida, Polychaeta) of the Arctic Ocean. Sarsia, 82, 203 - 236.","Moen, T. L. (2006) A translation of Bishop Gunnerus' description of the species Hydroides norvegicus with comments on his Serpula triqvetra. Scientia Marina, 70 S 3, 115 - 123. http: // dx. doi. org / 10.3989 / scimar. 2006.70 s 3115","Ben-Eliahu, M. N. & Hove, H. A. ten (2011) Serpulidae (Annelida: Polychaeta) from the Suez Canal-From a Lessepsian migration perspective (a monograph). Zootaxa, 2848, 1 - 147.","Nedved, B. T. & Hadfield, M. G. (2008) Hydroides elegans (Annelida: Polychaeta): A model for biofouling research. In: Flemming, H. - C., Murthy, P. S., Venkatesan, R. & Cooksey, K. E. (Eds.), Marine and Industrial Biofouling, Series in Biofuling, Springer-Verlag Berlin Heidelberg, 4, pp. 203 - 217. http: // dx. doi. org / 10.1007 / 978 - 3 - 540 - 69796 - 1 _ 11","Johnson, L. T. & Gonzalez, J. A. (2004) Staying afloat with nontoxic antifouling strategies for boats. California Sea Grant College Program Report Number T- 054. University of California, Oakland, California, 21 pp.","Johnson, L., Gonzalez, J., Alvarez, C., Takada, M. & Himes, A. (2006) Managing hull-borne invasive species and coastal water quality for California and Baja California boats kept in saltwater. California Sea Grant College Program Report Number T- 061. University of California, Oakland, California, 149 pp."]}

Copper (Cu) contamination is a potential threat to the marine environment due to the use of Cu-based antifouling paints. Cu stress on larval settlement of the polychaete Hydroides elegans was investigated, and this was linked to Cu stress on biofilms and on the biofilm development process. The inductiveness of young biofilms was more easily altered by Cu stress than that of old biofilms, indicating the relative vulnerability of young biofilms. This might result from changes in bacterial survival, the bacterial community composition and the chemical profiles of young biofilms. Cu also affected biofilm development and the chemical high performance liquid chromatograph fingerprint profile. The results indicate that Cu affected larval settlement mainly through its effect on the process of biofilm development in the marine environment, and the chemical profile was crucial to biofilm inductiveness. It is strongly recommended that the effects of environmentally toxic substances on biofilms are evaluated in ecotoxicity bioassays using larval settlement of invertebrates as the end point.

Hydroides elegans is a major fouling organism in tropical waters around the world, including Pearl Harbor, Hawaii. To determine the importance of initial surface characteristics on biofilm community composition and subsequent colonization by larvae of H. elegans, the settlement and recruitment of larvae to biofilmed surfaces with six different initial surface wettabilities were tested in Pearl Harbor. Biofilm community composition, as determined by a combined approach of denaturing gradient gel electrophoresis and fluorescence in situ hybridization, was similar across all surfaces, regardless of initial wettability, and all surfaces had distinct temporal shifts in community structure over a 10 day period. Larvae settled and recruited in higher numbers to surfaces with medium to low wettability in both May and August, and also to slides with high wettability in August. Pearl Harbor biofilm communities developed similarly on a range of surface wettabilities, and after 10 days in Pearl Harbor all surfaces were equally attractive to larvae of Hydroides elegans, regardless of initial surface properties.

Multi-species natural microbial biofilms have been identified as sources of cues to induce larval attachment of a broad range of sessile marine invertebrates. However, the chemical identities of the cues originating from these films have not been fully characterized. In this study, we isolated and characterized 2 compounds from natural biofilms that induced the larval attachment of the polychaete Hydroides elegans. Biofilms were developed on Petri dishes in seawater in the field for 6 d, collected from the Petri dishes using cotton buds, and extracted using a combination of polar and non-polar solvents. The non-polar fraction was inductive to larval attachment and was thus further fractionated using HPLC. Bioassay-guided fractionation was used to locate the active com- pounds that were then structurally characterized by using nuclear magnetic resonance (NMR) spec- troscopy. Two inductive compounds were identified, a hydrocarbon (6,9-heptadecadiene) and a fatty acid (12-octadecenoic acid), which are the first chemical cues inductive to larval attachment of marine invertebrates isolated from natural biofilms.

Many quantitative models on the evolution of divergent life histories in marine inverte- brates have focused on egg size with little empirical support for the assumption that significant heri- tability for egg size is present. Narrow-sense heritability (h 2 ) is the proportion of total phenotypic variance in egg size that is made up of additive genetic variance, and is a predictor of short-term response to selection. We estimated h 2 for the trait of egg size in the polychaete worm Hydroides ele- gans using artificial selection and by using a half-sibling breeding design. The cumulative realized heritability was calculated as 0.58 for egg size in a population of H. elegans in Pearl Harbor, Hawaii. Similarly, the half-sib breeding design gave an estimate for h 2 of 0.45 for the same population. This indicates that there is substantial potential for egg size to respond to varying selective pressures, at least in the direction of increase.

The uptake of anthropogenic CO2 emissions by oceans has started decreasing pH and carbonate ion concentrations of seawater, a process called ocean acidification (OA). Occurring over centuries and many generations, evolutionary adaptation and epigenetic transfer will change species responses to OA over time. Trans-generational responses, via genetic selection or trans-generational phenotypic plasticity, differ depending on species and exposure time as well as differences between individuals such as gender. Males and females differ in reproductive investment and egg producing females may have less energy available for OA stress responses. By crossing eggs and sperm from the calcareous tubeworm Hydroides elegans (Haswell, 1883) raised in ambient (8.1) and low (7.8) pH environments, we observed that paternal and maternal low pH experience had opposite and additive effects on offspring. For example, when compared to offspring with both parents from ambient pH, growth rates of offspring of fathers or mothers raised in low pH were higher or lower respectively, but there was no difference when both parents were from low pH. Gender differences may result in different selection pressures for each gender. This may result in overestimates of species tolerance and missed opportunities of potentially insightful comparisons between individuals of the same species.

We report here the 6.0-Mb draft genome assembly of Pseudoalteromonas luteoviolacea strain HI1 using Roche 454 and PacBio single-molecule real-time hybrid-sequencing analysis. This strain is of biological importance since it has the capacity to induce the settlement and metamorphosis of the serpulid polychaete Hydroides elegans and the coral Pocillopora damicornis .

The serpulid tubeworm Hydroides sanctaecrucis is recorded from tropical northern Australia for the first time. This species is native to the Caribbean, where it is common in coastal lagoons, and has not previously been reliably recorded from the western Pacific. The species was identified after heavy and unusual serpulid fouling was noted on vessel hulls in Cairns, north Queensland, and hull fouling is considered the most probable vector for its translocation to the western Pacific. In northern Australia, H. sanctaecrucis displays similar traits to other fouling and now widely dispersed fouling serpulids such as Hydroides elegans, H. ezoensis and Ficopomatus enigmaticus.

This study investigated the effect of UV-A and B (UVR) on larval settlement of the poly- chaete Hydroides elegans through their influence on biofilms. Multispecies biofilms were treated with 3 doses of UVR (10, 30 and 80 KJ m -2 ) at 2 environmentally realistic irradiance levels (4 and 2 W m -2 ) in the laboratory, and their ability to induce the settlement of H. elegans larvae was then exam- ined in both laboratory and field conditions. For the field experiment, only 10 and 80 KJ m -2 doses under 4 W m -2 were used. In addition, this study evaluated the effects of UVR on monospecies bacte- rial biofilms and then their ability to induce larval settlement in the laboratory. Results demonstrated that the ability of multispecies biofilms to trigger larval settlement could be compromised as a result of enhanced UVR exposure. Although larval settlement on multispecies biofilms treated with the low- est UV dose (at both irradiance levels) was at the same level as that of the control, the exposure of biofilms to the highest dose significantly reduced their larval settlement triggering ability. Further- more, UVR treatments decreased the percentage of metabolically active bacterial cells in mono- species biofilms; the effect increased with increasing UV dose. Larval settlement response to mono- species biofilms decreased with increasing UV dose, suggesting that the bacterial metabolic activity in biofilms is essential for the biofilms to have an inductive effect on larval settlement. This study sug- gests that enhanced UVR, which might occur due to ozone depletion, may have a significant effect on the larval settlement of H. elegans by affecting a biofilm's inductive cues.

In the biogeographical and taxonomical literature before the 1980s there was a wide perception that widespread, often referred to as ‘cosmopolitan’, species were very common among polychaetes. Here we discuss the origins of this perception, how it became challenged, and our current understanding of marine annelid distributions today. We comment on the presence of widely distributed species in the deep sea and on artificially extended ranges of invasive species that have been dispersed by anthropogenic means. We also suggest the measures needed to revolve the status of species with reported cosmopolitan distributions and stress the value of museum collections and vouchers to be associated with DNA sequences in resolving species distributions.

The polychaete, Hydroides elegans, is a tube-building worm that is widely distributed in tropical and subtropical seas. It is a dominant fouling species and thus a major target organism in antifouling research. Here, the first high-throughput proteomic profiling of pre-competent and competent larvae of H. elegans is reported with the identification of 1,519 and 1,322 proteins, respectively. These proteins were associated with a variety of biological processes. However, a large proportion was involved in energy metabolism, redox homeostasis, and microtubule-based processes. A comparative analysis revealed 21 proteins that were differentially regulated in larvae approaching competency.