Eualetes tulipa

Pictures

Picture	Title	Caption	Copyright
Title Subtidal specimens Caption Subtidal specimens of Eualetes tulipa attached to the walls of the Planta Centro power plant channel in Venezuela. Only the apertures of the organisms and the tip of the feeding tube are visible, as the rest of the shell is covered by epibionts. Copyright Ana Karinna Carbonini	Subtidal specimens	Subtidal specimens of Eualetes tulipa attached to the walls of the Planta Centro power plant channel in Venezuela. Only the apertures of the organisms and the tip of the feeding tube are visible, as the rest of the shell is covered by epibionts.	Ana Karinna Carbonini

Identity

Preferred Scientific Name

• Eualetes tulipa (Chenu, 1843)

Other Scientific Names

• Tripsycha (Eualetes) tulipa Chenu, 1843
• Vermetus alii
• Vermetus sp. Cuvier, 1800

Summary of Invasiveness

E. tulipa is a vermetid gastropod that inhabits shallow waters in warm temperate and tropical seas. The shell measures about 5 cm in length and 3 cm in width. Adults are attached to hard substrates, usually to artificial structures, however, it has also been reported in reefs and mangrove roots. Although the adults are rather more solitary than colonial, populations may form clusters with densities of up to 100 individuals/m². In the southern Caribbean, this species has been reported to reproduce throughout the year, the females brooding up to 54 capsules in the mantle cavity, each containing about 280 eggs of which around 190 hatch as veliger larvae that settle within 24 hours after hatching. This species has been reported in Panama Bay (tropical Pacific), in the Venezuelan Caribbean and at several localities in Hawaii, where it has been reported as an invasive species since 1997. Knowledge on the distribution of this species is very scarce. If indeed this is an invasive rather than a cryptogenic species either in the Caribbean or in the Hawaiian Islands, invasion is probably via vessel biofouling.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Mollusca
•             Class: Gastropoda
•                 Subclass: Caenogastropoda
•                     Order: Littorinimorpha
•                         Superfamily: Vermetoidea
•                             Family: Vermetidae
•                                 Genus: Eualetes
•                                     Species: Eualetes tulipa

Notes on Taxonomy and Nomenclature

The Vermetidae have one of the most complicated histories in taxonomy, mainly due to the fact that the first taxonomic insights assumed many plastic characters of the morphology were apomorphies (Keen, 1961; Morton, 1965; Keen, 1982; Bieler, 1989; Bieler, 1996).

The first attempt to classify species of this family was carried out by Mörch in the 1860s, who included within the same group, species from other families belonging to different taxa (Keen, 1961; Bieler, 1996). A decade later, Keen (1961) proposed a new classification recognizing 5 major genera: Dendropoma, Petaloconchus, Tripsycha, Vermetus and Serpulorbis. This taxonomic scheme was supported by Morton (1965) who also established the phylogenetic relationships and placed them along with other vermiform snails within the Superfamily Cerithioidea. The variables used to classify species under this scheme were mostly the morphology of both the protoconch and the teleoconch, the operculum, the spiralization, and some anatomical features.

Although this classification represented a significant advance in the taxonomy of the group, more research showed that many species did not fit well within it (Hadfield et al., 1972; Bieler, 1995; Schiaparelli, 1996; Schiaparelli and Cattaneo-Vietti, 1999), which questioned the validity of using morphological characters to define genera and species (Gould, 1994; Bieler, 1996; Schiaparelli and Métivier, 2000). On the other hand, several authors showed that a more accurate phylogeny of the family did not correspond with the phylogeny proposed by Morton (Hughes, 1993; Gould, 1994; Healy, 1988; Lydeard et al., 2002). At present, the family is still under taxonomic revision. Eualetes tulipa was originally described as Vermetus tulipa by Chenu (1842-1853), and Keen (1971) renamed this species as Trypsicha (Eualetes) tulipa.

Description

E. tulipa is a marine gastropod species of the Vermetidae family, commonly known as "worm gastropods" due to the tube shape of their shells. Vermetids are sessile and are either attached or buried on hard substrates. Once the juvenile attaches to the substrate, the adult shell grows by coiling itself at a right angle to the nuclear whorls of the juvenile shell (Keen, 1971). This produces a large morphological plasticity in the adults (Hadfield et al., 1972; Safriel and Hadfield, 1988; Gould, 1994; Bieler, 1995; Schiaparelli, 1996; Schiaparelli and Cattaneo-Vietti, 1999).

Distribution

In the original description of E. tulipa made by Chenu (1842-1853) as Vermetus tulipa, no locality was specified for the collected material. Keen (1971) reported this species at several localities in Panama Bay, including the boulder reef at Verra Cruz (Hughes, 1985), and Vega and González (2002) reported it at Veraguas Province, both in the east Pacific Ocean. In Hawaii, E. tulipa has been reported as a non-indigenous, introduced species in 32 localities including Kane‘ohe Bay, Hilo Harbour, O’ahu Waikiki and O’ahu Honolulu since at least 1997. In these Hawaiian localities, individuals are mainly found attached to artificial substrates such as concrete pilings and walls, metal and wood substrates but also in natural environments such as reefs and mangrove roots (Coles and Eldredge, 2002; Coles et al., 2002; Schluker, 2003; Ray, 2005; database from the Bishop Museum, Hawaii). In the Venezuelan Caribbean, this species (reported as Vermetus sp.) has been observed since at least 1986 in the central west coast nearby Puerto Cabello colonizing the walls of the intake-cooling sea water channel of a thermoelectric power plant (Miloslavich and Penchaszadeh, 1992). This species has also been observed at other two localities within the area colonizing other artificial substrates: the first is on the concrete pilings at the deck of El Palito oil refinery and the other is on the metal remains of a ship wreckage (“Sesostris”) from World War II at Isla Larga (Miloslavich and Penchaszadeh, 1992; Osman and Shirley, 2007).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

Knowledge on the distribution of this species is very scarce, so at this moment there is insufficient information to establish how it has spread between the Caribbean and the Pacific. If indeed this is an invasive rather than a cryptogenic species (defined by Carlton, 1996 as a species that cannot be reliable demonstrated either as being introduced or endemic), the species is probably carried via fouling of vessels. The most common navigational route between the tropical Pacific and the Caribbean is through the Panama Canal, a bridge of freshwater connecting the two oceans. It takes approximately 8 to 10 hours for a ship to cross this canal although ships may have to queue for up to a few days in relatively estuarine conditions since each time a ship transits the canal. Miloslavich (Universidad Simón Bolivar, Venezuela, personal communication, 2009) found that adults of this species can survive at least 3 days in freshwater (0º/oo) by sealing their operculum, and can continue with normal activities after placed again in seawater. The density of the population at the Planta Centro power plant channel (Venezuelan Caribbean) was about 30 individuals/m² in 1986 (Miloslavich and Penchaszadeh, 1992), however this density has increased significantly in the last two decades to 53 and 72 individuals/m² respectively for 2006 and 2009.

Risk of Introduction

No information is available on risk of introduction, however, this species might be introduced either as adults attached to the hull or other ship surfaces, or as larvae in ballast waters. The former is more likely as adults of E. tulipa have been shown to tolerate immersion in freshwater for several days. The latter is more unlikely as the veliger larvae usually settle within 24 hours after hatching from the egg capsules (Miloslavich and Penchaszadeh, 1992).

In general,some species of vermetids are considered to have great dispersive and even invasive potential (Bieler, 1996; Strathmann and Strathmann, 2006). Although surveys evaluating their dispersion capacities, competitive abilities and propagule viability have not been carried out, vermetids are known for their high tolerance to adverse conditions, such as lack of food, clean water, obstacles (Schiaparelli and Cattaneo-Vietti, 1999; Strathmann and Strathmann, 2006), their ability to attach to different artificial substrates (Miloslavich and Penchaszadeh, 1992; Schiaparelli et al., 2002) and to have extra-embryonic food sources that allow the progeny to spend large periods drifting (Strathmann and Strathmann, 2006).

Habitat

E. tulipa lives in tropical shallow waters, from the intertidal to the subtidal zone. In the Venezuelan Caribbean, it has been reported to live between 0 and 5 m depth, with significantly higher densities (up to 30 individuals/m²) in the first 3 m of depth (Miloslavich and Penchaszadeh, 1992). It is frequently associated with artificial hard substrates such as concrete, iron or wood pilings, floating docks, concrete walls, flat blocks, and the iron hulls of wrecked ships. In Hawaii, it has also been reported to be associated with coral reefs and mangrove roots (database from the Bishop Museum, Hawaii).

Habitat List

Biology and Ecology

Genetics

The complete mitochondrial genome of E. tulipa has been sequenced and published, see Rawlings et al. (2010).

Reproductive Biology

E. tulipa is a dioecious species that reproduces throughout the year. Females brood up to 54 spherical egg capsules in the mantle cavity. These capsules measure about 2.3 mm in diameter and contain about 290 eggs each measuring 240 mm in diameter. Only about 188 eggs develop to hatch as veliger larvae, the rest are consumed by the developing embryos as nurse eggs. Near the time of hatching, the capsule increases its volume and measures about 3.4 mm in diameter. Larval size at hatching is about 454 mm, they show positive phototaxis and settle within 24 hours. Once settled, the juvenile resorbs the velum and begins growing the “tube” like shell. Developmental time from the egg to the hatching stage is 21 days (Miloslavich and Penchaszadeh, 1992). The biochemical content (protein, glycogen and lipid) of the eggs and hatchlings of E. tulipa was measured by Miloslavich (1996). Each egg contain near 2 mg of protein, 15 mg of glycogen, and 7 ug of lipid. At hatching, each veliger larva contains about 0.7 mg of protein, 24.6 mg of glycogen, and 9.8 mg of lipid. Eggs are mostly constituted by high molecular weight proteins (more than 100 Kd), while hatchlings are constituted by several low molecular weight proteins and some high molecular weight proteins.

There is no data available on the growth of this species, however, some tube regeneration experiments have been conducted both in the field (at the Planta Centro power plant in Venezuela) and under laboratory conditions. Tubes grow about 1.1 mm/week in the field and 2.5 mm/week under laboratory conditions (Miloslavich et al., unpublished data, P Miloslavich, Universidad Simón Bolivar, Venezuela, personal communication, 2009). Schiaparelli (1996) noted that when animals of this family are kept in an aquarium, with little water movement, they produce feeding-tubes probably to attain the best employment of gills and/or mucus web.

E. tulipa, like most vermetids, is a suspension-feeder that feeds continuously by means of a mucous thread that is extended by pedal tentacles up to 15 cm in length from the feeding tube, taking advantage of organic suspended matter brought by the water currents. This web is hauled and ingested at intervals of about 130 s depending on the individual and the environmental conditions. As adults live close to each other, when one of them hauls its web, it inevitably pulls some of its neighbors web, there is no evidence of synchronicity in this activity between them (Hughes, 1985).

As a generally tropical hard substrate species, E. tulipa is found in a very diverse community. At the Planta Centro power plant in Venezuela, the fouling community in the intertidal zone is dominated by two vermetids of the genus Petaloconchus (Petaloconchus varians, and Petaloconchus sp., an unidentified orange species) as well as barnacles (genus Balanus and Cthamalus) and oysters (Isognomon alatus). The subtidal zone contains tunicates, either solitary or colonial, bryozoans, cirripedes, anemones, polychaetes (sabellids, serpulids, and spirorbids), hydrozoans, octocorals, sponges, and algae. Other associated fauna are echinoderms (Diadema antillarum, Echinometra lucunter, Isostichopus badionotus), other gastropods (species of the genus Littorina and Thais haemastoma), crabs, amphipods, and fish (Losada et al., 1988). In Panama, Keen (1971) reported this species to be associated with Stephoma pennatum, a vermicularid gastropod.

E. tulipa has been reported to live at temperatures varying between 26 and 30ºC and salinities between 34 and 36º/oo (Hughes, 1985; Miloslavich and Penchaszadeh, 1992). The concentration of organic suspended matter measured at one of the population sites (Planta Centro power plant channel, Venezuela) varied between 1.7 to 21.8 mg/L depending on the season (higher values in the rainy season and lower values in the dry season), being the annual mean of 6.2 mg/L. At this site, the power plant machinery creates a continuous suction current of about 0.2 m/sec, from the sea to the inside of the plant, all through the channel. This flow of water guarantees a continuous supply of organic suspended matter for the vermetids. At a nearby population (Isla Larga), the conditions are similar with the exception of the current. This population is not nearly as dense as that from the power plant channel, the individuals being rather solitary. It is most likely that an increase in the amount of organic suspended matter in the water may lead to an increment in population size.

Water Tolerances

Natural Enemies

Notes on Natural Enemies

Field observations in the Planta Centro power plant channel (Venezuela / Caribbean) have shown the presence of the predator gastropod Thais haemastoma living among individuals of E. tulipa. Empty tubes of E. tulipa are used by fish of the family Blenidae (Miloslavich, 1987).

Means of Movement and Dispersal

Adults are sessile throughout their life. The only stage with dispersal potential is the veliger larvae which has a short planktonic life of about 24 hours.

Pathway Vectors

Environmental Impact

Shells of E. tulipa increase spatial heterogeneity and create microhabitats. Also, the shells of this species either dead or alive are used by a variety of epifauna and turf macroalgae in the subtidal zone. In general, the only visible part of most individuals is the tube opening with the operculum, the rest being covered by other organisms such as sponges, hydrozoans and turf macroalgae (the radula prevents the establishment of other organisms inside the tube or blocking of the shell aperture). The establishment of large epifauna associated with the vermetids can change the water flow and circulation pattern near them, facilitating larval settlement (Losada et al., 1988). Vermetids are also known to form solid organic reefs at sea level and have been used as highly reliable biological sea-level indicators for palaeo sea-level reconstruction (Laborel, 1986; Laborel and Laborel-Deguen, 1996; Morhange et al., 1998; Antonioli et al., 1999). Nevertheless, Schiaparelli et al. (2006) showed that the degree of habitat plasticity of Dendropoma petraeum, a Mediterranean species commonly used for these purposes, along with the almost total lack of knowledge on the precision of other species, leads to conclude that a common value of precision cannot be maintained.

Impact on Biodiversity

The impact of E. tulipa on the local biodiversity has not been studied; however, as mentioned earlier, this species is associated with a very diverse hard substrate community that benefits from the spatial heterogeneity created by the vermetids.

Risk and Impact Factors

Impact mechanisms

• Competition
• Filtration
• Fouling
• Rapid growth

Impact outcomes

• Ecosystem change/ habitat alteration
• Modification of natural benthic communities

Invasiveness

• Abundant in its native range
• Fast growing
• Gregarious
• Has high reproductive potential
• Proved invasive outside its native range
• Tolerant of shade

Likelihood of entry/control

• Difficult to identify/detect as a commodity contaminant
• Difficult to identify/detect in the field
• Highly likely to be transported internationally accidentally

Similarities to Other Species/Conditions

There are approximately 250 vermetid species worldwide (Hadfield, 1970), however, as mentioned earlier, vermetid taxonomy is complicated (Keen, 1961; Bieler, 1996) and few genera are defined by valid autopomorphic characters (Gould, 1994). Schiaparelli et al. (2006) stated that the complex taxonomy of this family has on occasion led researchers to mix different species, even belonging to different genera under a single name, and synonymies in this group are very common. With the recent development of molecular techniques, the taxonomy of vermetid species may be solved. Using these methods, Calvo et al. (2009) discovered that there are at least four different cryptic species in what was called the Dendropoma petraeum complex from 18 different localities in the Mediterranean Sea. Also with molecular techniques, Faucci et al. (2007) established the connectivity between vermetid populations of the nine species that are distributed throughout the Hawaiian Archipielago, proposing that species with limited larval dispersal potential show structured populations suggesting little or very limited connectivity among islands. In the intertidal zone of the Planta Centro power plant channel, E. tulipa cohabits with two other vermetid species of the genus Petaloconchus, their shells forming dense multispecies colonies (Weinberger, 2008).

Keen (1971) reports another similar species of the genus Eualetes (E. centiquadra) which is distributed from the Gulf of California to southern Mexico in the Pacific Ocean. Within the vermetid family, the most similar species to E. tulipa are those of the genus Vermetus. In the OBIS database (Ocean Biogeographic Information System), there is no record of E. tulipa; however, there are ten species of Vermetus (V. africanus, V. alli, V. corallinaceus, V. corrugatus, V. erectus, V. formosus, V. granulatus, V. natalensis, V. periscopium and V. quincunx), and many others have been described (e.g. Chenu (1842-1853)). Until the taxonomy of the family is clarified, it is uncertain if these species reflect the actual biodiversity of the genus.

Prevention and Control

Control

Physical/mechanical control

There are no reports of physical or mechanical control of this species; however, when the Planta Centro power plant turbines undergo maintenance, all biofouling organisms are mechanically removed (personal communication with the office of Hygiene and Industrial Safety of the Planta Centro power plant, Venezuela from P Miloslavich, Universidad Simón Bolivar, Venezuela, personal communication, 2009).

Chemical control

There are no reports of chemical control of this species; however, the Planta Centro power plant turbines usually receive chlorine applications to prevent the establishment of biofouling (personal communication with the office of Hygiene and Industrial Safety of the Planta Centro power plant, Venezuela from P Miloslavich, Universidad Simón Bolivar, Venezuela, personal communication, 2009).

Gaps in Knowledge/Research Needs

Work is needed in several areas, firstly in the area of molecular biology, in order to verify that all the populations reported as E. tulipa are indeed the same species. Secondly on population dynamics and the effect that an increase in organic suspended matter in the seawater might have on population size. Thirdly on the effect that the species may have on the community (at different population densities), including increase in spatial heterogeneity and ecological role.

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Links to Websites

Contributors

30/06/09 Original text by:

Patricia Miloslavich, Universidad Simón Bolívar, Venezuela

Distribution Maps

• = Present, no further details
• = Evidence of pathogen
• = Widespread
• = Last reported
• = Localised
• = Presence unconfirmed
• = Confined and subject to quarantine
• = See regional map for distribution within the country
• = Occasional or few reports

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