Hydroides elegans

Invasion History

First Non-native North American Tidal Record: 2001
First Non-native West Coast Tidal Record: 2001
First Non-native East/Gulf Coast Tidal Record: 2011

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General Invasion History:

Hydroides elegans is a serpulid tubeworm, originally described from Port Jackson, Sydney, Australia, which is now widely distributed in in tropical-to-warm temperate marine waters (Zibrowius 1971; ten Hove 1974; Keough, and Ross 1999; Bastida-Zavala et al. 2002). Its origin is unknown but is usually presumed to be somewhere in the Indo-Pacific. Molecular studies indicate a high degree of similarity among global populations, probably due to extensive genetic exchange from ship transport (Pettengill et al. 2007). However, much of this transport occurred before serpulids were carefully identified. The known global distribution of this tubeworm is additionally complicated by the fact that it was long confused with the northeast Atlantic species Hydroides norvegica (Zibrowius 1971). Genetic and morphological studies may clarify the distribution and origin of this tubeworm. Currently, H. elegans is considered cryptogenic in most of its North American waters (Bastida-Zavala et al. 2021), except for areas north of its established range, which have recently been colonized (e g. San Francisco Bay, Humboldt Bay, and a failed population in Woods Hole, Massachusetts). 

Hydroides elegans frequently has been considered introduced on both sides of the Atlantic, the Mediterranean Sea, the northeast Pacific (Mexico-California), Hawaii, Russia and Japan, and New Zealand. In most localities, it is restricted to, or concentrated in, polluted harbors (Zibrowius 1971; Carlton 1979; Bastida-Zavala 2008). Since its native range is unknown, Bastida-Zavala et al. (2017) have considered it cryptogenic in US waters. We treat it here as cryptogenic in Hawaii, Mexico, southern California, the Caribbean, and the Gulf of Mexico. However, there are recent range extensions, and probable introductions to Morro and San Francisco Bays (Blum et al. 2007; Needles and Wendt 2012), and a non-established occurrences in Massachusetts and Humboldt Bay (Bastida-Zavala et al. 2017). 

North American Invasion History:

Invasion History on the West Coast:

On the west coast of North America, Hydroides elegans was first reported from San Francisco Bay, at Mare Island Naval Station, on the hull of the submarine U.S.S. 'Narwhal' (in 1929, Carlton 1979). Established populations were first found in 1931 in Los Angeles-Long Beach Harbors (Zibrowius 1970, cited by Carlton 1979). In southern California, H. elegans was collected from San Diego Bay (1st record 1948) and Marina del Rey (in 1972), and is locally abundant (Carlton 1979; Bastida-Zavala 2008, Ruiz et al., unpublished data). This tubeworm was also collected in the Gulf of California at Bahia de La Paz, Mexico in 1991, where it was abundant (Bastida-Zavala 2008). The southward extent of its range in the Eastern Pacific is not known. In 2001, H. elegans was found on settling plates in Richmond Marina, San Francisco Bay (Blum et al. 2007), extending the northward range of this species in the Eastern Pacific. Live specimens have been found in ship fouling, in Vancouver, British Columbia (Sylvester et al. 2011), but are unlikely to survive that far north.

Invasion History on the East Coast:

Hydroides elegans was found on a boat in 'southern Florida' in 1951 (identified as H. norvegica, Hartman 1952). There are scattered records from Biscayne Bay and Indian River Lagoon (Walters 2001; 2004, Ruiz et al., unpublished data). In 2011, this subtropical worm was surprisingly discovered in Eel Pond, Woods Hole, Massachusetts, in November and December.  It developed extensive colonies and survived the winter of 2011–2012 (ICES Advisory Committee on the Marine Environment 2012; James T. Carlton, personal communications, 2011-2013).

Invasion History on the Gulf Coast:

Hydroides elegans (as H. norvegica) was found on the hull of a boat in Corpus Christi Bay, Texas (Hartman 1952). A specimen was collected from Safe Harbor, Key West, Florida (in 1970, USNM 45242, U.S. National Museum of Natural History 2007). This worm was collected on fouling plates in Tampa Bay in 2002 (Ruiz et al. unpublished data). In Mexican waters, H. elegans was collected in Veracruz, Mexico in 1960 and 1996, and in Campeche in 1999 (Bastida-Zavala and ten Hove 2002).

Invasion History in Hawaii:

Hydroides elegans was first collected in Pearl Harbor, Oahu in 1929 (Straughan 1969 as H. norvegica; Coles et al. 1999) and subsequently found in Kanaohe Bay (1935, Straughan 1969) and Pearl Harbor (1929, Straughan 1969, Coles et al. 1999). Hydroides elegans has usually been regarded as introduced in Hawaii (Carlton and Eldredge 2009), but this could be part of a broader Indo-Pacific distribution (Bastida-Zavala et al. 2017). 

 

Invasion History Elsewhere in the World:

The earliest known record of Hydroides elegans is from Italy in 1844 (as Eupomatus pectinatus Philippi, 1844), but this could represent a very early introduction (Bastida-Zavala et al. 2017). Another early record was from the Virgin Islands, in the Caribbean (Krøyer in Mörch, 1863, as H. abbreviata). In 1883, it was described as Eupomatus elegans from Port Jackson, Australia, by Haskell in 1883 (Bastida-Zavala et al. 2017). Hydroides elegans are often considered of Indo-Pacific origin, but the other early records make it difficult to assign a native region. 
 
We are treating most of the records of Hydroides elegans in tropical, subtropical and warm-temperate regions as cryptogenic. It is established in the south Atlantic, in Guanabara Bay, Rio de Janeiro, Brazil and on the coast of Africa from Senegal (1954, Sourie 1954; Fauvel and Rullier 1957, both cited by Zibrowius 1971), Ghana, (Zibrowius 1971), Luanda, Angola (Kirkegard 1959, cited by Zibrowius 1971), and Cape Town, South Africa (Zibrowius 1971). Occurrences in Mar del Plata, Argentina probably represent an introduction (Orensanz et al. 2002). It is established in the Azores (Morton and Britton 2000). As noted above, H. elegans was discovered very early in the Mediterranean (Philippi 1844, cited by Bastida-Zavala et al. 2017), and collected in 1888 in the Bay of Naples (Lo Bianco 1893, cited by Zibrowius 1971). It now occurs in harbors throughout the Mediterranean, from France and Spain to Israel, Egypt, and Turkey (Zibrowius 1971; Cornelio and Manzoni 1999; Pancuci-Papadopoulou et al. 2005; Cinar 2006; Galil 2007; US National Museum of Natural History 2008). In England and the Netherlands, H. elegans has occurred only in thermal effluents and heated docks (ten Hove 1974: Zibrowius and Thorp 1989). 
 
Hydroides elegans is widely distributed in the Indian Ocean and West Pacific, from Mozambique, the Red Sea, and the Persian Gulf, to Indonesia, the Philippines, Australia, China, and Polynesia (Zibrowius 1971; Rajagopal et al. 1997; Keough and Ross 1999; Wang and Huang 1999; US National Museum of Natural History 2008) and may be of Indo-Pacific origin (Zibrowius 1971; Pettengill et al. 2007). However, it occurs in New Zealand (first recorded in 1952, Cranfield et al. 1998), Japan (first record 1932, Asakura 1992), and the Vladivostok area of Russia (2000, Bagaveeva and Zvyagintsev 2000; Zvyagintsev 2003), where it is confined to thermal effluents. 

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Description

Hydroides elegans secretes a calcareous tube, as do other serpulid polychaetes. Serpulids have a feathery crown of modified prostomial palps, called radioles (the prostomium is the first segment, projecting above the mouth). The radioles can be folded and withdrawn into the tube. One of the radioles is modified to form an operculum, which acts as a plug when the animal contracts. The peristomium (segment behind the mouth) is folded back to form a collar, which bears uniramous parapodia, with a distinctive set of collar chaetae, with spines or serrations. The collar is the first of seven thoracic chaeta-bearing segments (chaetigers). The subsequent segments have biramous parapodia. The dorsal branch of the parapodium is called the notopodium; the ventral is the neuropodium. Chaetae in the two branches and along the body can vary greatly in their morphology, which can be critical in the taxonomy of serpulids. Description from: Barnes 1983; Bastida-Zavala and ten Hove 2002). 
 
The tube of H. elegans is white, and 1.3–2.5 mm in diameter. The tubes are fragile and variable, sometimes having transverse ridges and occasionally having longitudinal ridges, but are usually smooth. The tubes lack peristomes (flared openings). The tubes are usually flattened on the dorsal (upper) surface. The branchial (gill) crown consists of about 10 radioles each on the left and right sides of the mouth. It comprises about 1/4 of the worm's length. The operculum is roughly funnel-shaped, with 23–24 radii, each with rounded tips, a concave distal surface, and a circular row of 13–15 terminal spines, usually barbed with up to 4 spinules. The peduncle is cylindrical. The grooves between the radii are usually about 1/3 of the funnel length. The spines are longer than the radii, and T-shaped, with expanded tips, and have a single spinule at the base. The verticil (ring of distal spines) may have or lack a central tooth. The thorax consists of 7 segments. There are two kinds of collar chaetae: (1) thicker bayonet chaetae, with two teeth at their base, a reap of denticles below the teeth, and fine, saw-like denticles on the distal edge; (2) hair-like (capillary) chaetae. The subsequent thoracic segments bear short, rasp-like setae, called uncinae, and limbate chaetae. The abdomen has about 41 segments (35–57, n=5). The overall length is about 8.5 mm (5–13, n=5). The worm is yellow to light brown. (Description from Bastida-Zavala and ten Hove 2002; Cinar 2006; Ben-Eliahu and ten Hove 2011). 
 
Historically, the cosmopolitan tropical-subtropical harbor serpulid, H. elegans was frequently misidentified as H. norvegica, a Northeast Atlantic species associated with clear, cold waters, found from Norway to Morocco. Zibrowius (1971) clarified distinctions between these species. The collar chaetae in H. elegans have a saw-like row of denticles on their distal portion. Additional features distinguish the two species. Early records of H. norvegica (before the 1970s) from tropical or subtropical areas usually refer to H. elegans (Zibrowius 1971; ten Hove 1974; Bastida-Zavala and ten Hove 2002; Ben-Eliahu and ten Hove 2011). 

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Taxonomy

Ecology

General:

Life History – The serpulid polychaete Hydroides elegans feeds by extending its feathery gills and trapping plankton in the water column, which are transported by cilia to the mouth. The sexes are separate, as in most serpulid species. The larvae are planktotrophic and spend about 4-8 days in the plankton at 15–30 °C, with development being slower at low temperatures and low food levels (Qiu and Qian 1997). 
 
Ecology – Hydroides elegans tolerates salinities as low as 15 PSU, and successfully reproduces at 20 PSU (Qiu and Qian 1997) but is usually associated with marine salinities of 30-37 PSU (Bastida-Zavala and ten Hove 2002). However, in the Suez Canal, it was found in the Great Bitter Lakes and Lake Timsah, where salinities exceeded 40 PSU. To our knowledge, the temperature tolerance of H. elegans has not been studied experimentally. Hydroides elegans is comparatively tolerant of tributyltin antifouling compounds, wood preservative chemicals, low oxygen, and hydrogen sulfide, and benefits from the dense phytoplankton concentrations in polluted harbors (Udhayakumar and Karande 1996; Tarakanadha et al. 2004). It secretes a calcareous tube, often irregularly coiled, on hard surfaces such as rocks, pilings, floats, shells, corals, mangroves, and ships’ hulls. While it often forms dense aggregations on surfaces, it is not known to form reefs. The aggregations of worms appear to result from hydrodynamic processes and passive settlement, rather than by chemical cues and active swimming (Walters et al. 1999). 

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	None
General Habitat	Mangroves	None
General Habitat	Grass Bed	None
General Habitat	Canals	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Salinity (‰)	15	Laboratory observations, Qiu and Qian 1997
Minimum Reproductive Temperature	15	Laboratory observations, Qiu and Qian 1997
Maximum Reproductive Temperature	30	Laboratory observations, Qiu and Qian 1997
Minimum Reproductive Salinity	20	Laboratory observations, Qiu and Qian 1997
Minimum Duration	4	Larval period, 30 C, 35 ppt, high food- Qiu and Qian 1997
Maximum Duration	8	Larval period, 15 C, 20 ppt, low food- Qiu and Qian 1997
Maximum Length (mm)	33	Zibrowius 1971
Broad Temperature Range	None	Warm temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Economic impacts- Hydroides elegans (mentioned as H. norvegica) is well-known as a ship-fouling organism and is the most common annelid reported on ships' hulls (Woods Hole Oceanographic Institution 1952; Nelson-Smith 1971). Their tubes contribute less drag than the shells of barnacles, so they may have less effect on ships' speed, but that also means that they are less likely to be dislodged by current (Nelson-Smith 1971). This polychaete was a dominant fouling organism in polluted marinas in the Aegean Sea, Turkey, but was rare in clean harbors (Kocak and Kucuksezgin 2000). H. elegans is comparatively tolerant of tributyltin antifouling compounds, wood preservative chemicals, low oxygen, and hydrogen sulfide, and benefits from the dense phytoplankton concentrations in polluted harbors (Udhayakumar and Karande 1996; Tarakanadha et al. 2004). Hydroides elegans was reported to create fouling problems in industrial water systems in Italy (Parenzan 1965, Paoletti and Sebastao 1973, cited by Zibrowius 1991). Heavy settlement of H. elegans in oyster beds in Hiroshima Bay, Japan reduced oyster settlement and caused extensive mortality (Miura and Kajihara 1984, cited by Hewitt et al. 2008). 
 
Ecological Impacts- Hydroides elegans is an abundant fouling organism in many warm-water harbors and is a probable competitor with other fouling organisms This polychaete was a dominant fouling organism in polluted marinas in the Aegean Sea, Turkey, but was rare in clean harbors (Kocak and Kucuksezgin 2000). It was also one of the dominant organisms on both fixed and moving artificial structures in Sydney Harbor, Australia (Glasby et al. 2000). Heavy settlement of H. elegans in oyster beds in Hiroshima Bay, Japan reduced oyster settlement and caused extensive mortality (Miura and Kajihara 1984, cited by Hewitt et al. 2008). Tubeworms (Hydroides spp., probably including H. elegans were dominant fouling organisms on the shells of living and dead oysters in the Mosquito Lagoon, Florida (Walters 2001; Boudreaux et al. 2006), and are likely to compete with adult oysters for food and oyster spat for space. The extensive masses of calcareous tubes created by these worms are likely to create habitat for other organisms. 

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Regional Impacts

NWP-2	None		Economic Impact	Fisheries
Fouling of aquaculture cages (Wang and Huang 1993)
NWP-3b	None		Economic Impact	Fisheries
Heavy settlement of H. elegans in oyster beds in Hiroshima Bay, Japan reduced oyster settlement and caused extensive mortality (Miura and Kajihara 1984, cited by Hewitt et al. 2008).
MED-VI	None		Economic Impact	Shipping/Boating
This polychaete was a dominant fouling organism in polluted marinas in the Agean Sea, Turkey, but was rare in clean harbors (Kocak and Kucuksezgin 2000).
MED-VI	None		Ecological Impact	Competition
This polychaete was a dominant fouling organism in polluted marinas in the Agean Sea, Turkey, but was rare in clean harbors (Kocak and Kucuksezgin 2000).
NWP-3b	None		Ecological Impact	Competition
Heavy settlement of H. elegans in oyster beds in Hiroshima Bay, Japan reduced oyster settlement and caused extensive mortality (Miura and Kajihara 1984, cited by Hewitt et al. 2008).
NWP-3b	None		Ecological Impact	Habitat Change
Heavy settlement of H. elegans in oyster beds in Hiroshima Bay, Japan reduced oyster settlement and caused extensive mortality (Miura and Kajihara 1984, cited by Hewitt et al. 2008).
MED-VII	None		Economic Impact	Industry
Hydroides elegans was reported to create fouling problems in industrial water systems in Bari, Italy (Paoletti and Sebastao 1973, cited by Zibrowius 1991).
MED-IV	None		Economic Impact	Industry
Hydroides elegans was reported to create fouling problems in industrial water systems in the Mar Piccolo, Taranto, Italy (Parenzan 1965, cited by Zibrowius 1991).
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Economic Impact	Shipping/Boating
Hydroides elegans has been found in fouling of boats and marinas in the Gulf of Mexico and southern Florida (Hartman 1952; Renaud 1956; Ruiz et al., unpublished data) and probably has some impact in boat and ship fouling in those regions.
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Ecological Impact	Competition
Tubeworms (Hydroides spp., probably including H. elegans) were dominant fouling organisms on the shells of living and dead oysters in the Mosquito Lagoon, Florida (Walters 2001; Boudreaux et al. 2006), and are likely to compete with adult oysters for food and oyster spat for space.
AUS-X	None		Economic Impact	Shipping/Boating
None
AUS-X	None		Ecological Impact	Competition
Hydroides elegans was also one of the dominant organisms on both fixed and moving artifical structures in Sydney Harbor, Australia (Glasby et al. 2000).
CIO-I	None		Economic Impact	Shipping/Boating
H. elegans is comparatively tolerant of tributyltin antifouling compounds, wood preservative chemicals, low oxygen, and hydrogen sulfide, and benefits from the dense phytoplankton conentrations in polluted harbors (Udhayakumar and Karande 1996; Tarakanadha et al. 2004).
CIO-I	None		Ecological Impact	Competition
H. elegans is comparatively tolerant of tributyltin antifouling compounds, wood preservative chemicals, low oxygen, and hydrogen sulfide, and benefits from the dense phytoplankton conentrations in polluted harbors (Udhayakumar and Karande 1996; Tarakanadha et al. 2004).
NWP-2	None		Economic Impact	Shipping/Boating
'Structures submerged in eastern Hong Kong waters for several months during winter and spring can be coated by H. elegans with a thickness up to 3.8 cm and wet-weight reaching 12.5 kg m-2 (Wang & Huang 1993).' (Sun et al. 2012).

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