Ficopomatus enigmaticus ( Fauvel, 1923 )

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, European Journal of Taxonomy 344, pp. 1-76 : 17-19

Ficopomatus enigmaticus ( Fauvel, 1923)

Figs 2 E–H, 3

Mercierella enigmatica Fauvel, 1923: 424–430 , figs 1a–o (type locality: Caen Canal, Normandy, France; on stems of Phragmites ( Poaceae grass), submerged wood, stones and shells in brackish water; associated with the bivalve Congeria cochleata (Nyst, 1835) , now Mytilopsis leucophaeata (Conrad, 1831) , the amphipod Corophium volutator (Pallas, 1766) and the bryozoan Membranipora lacroixii (Audouin, 1826) , now Conopeum reticulum ( Linnaeus, 1767)) .

Mercierella enigmatica – Fauvel 1933: 190 –192 (Lake Merritt, Oakland, San Francisco Bay; on pilings, piers, docks and boat hull associated with the amphipod Corophium bonelli Milne Edwards, 1830 (actually Monocorophium insidiosum (Crawford, 1937) sensu James T. Carlton , pers. comm.), the hydrozoan Syncoryne Steenstrup, 1842 (actually Sarsia tubulosa (M. Sars, 1835) sensu James T. Carlton , pers. comm.), and Mytilus Linnaeus, 1758 ). — Hartman 1952: 64 (Rockport, Texas; fouling). — Rioja 1943: 547 —550 (Puerto Quequén, Argentina). — Straughan 1969: 234 (Ala Wai Canal, Kewalo Basin and Pearl Harbor, Oahu, Hawaii). — Bailey-Brock 1976: 73 (Oahu and Maui islands, Hawaii; associated with channels, fish farms and oysters).

Ficopomatus enigmaticus – ten Hove & Weerdenburg 1978: 114 –116, figs 2e–i, 3d–e, l–q, 4a–d, s, aa–bb, nn–vv, zz, 5c (revision of the genus; Netherlands, Tunisia, South Africa, Australia, Japan, Uruguay, Argentina and United States: San Francisco, Texas and Hawaii). — Hoagland & Turner 1980: 60 (Oyster Creek Nuclear Generating Station, Barnegat Bay, New Jersey; 0.8–2 m; occurs in low densities on wood plates in the mouth of creeks flowing to the bay). — Zibrowius 1992: 91 (discussion about its origin). — ten Hove & van den Hurk 1993: 42 –44, figs 5A–D, 6A–C (Lake Tunis reefs; intertidal to 1.9 m, salinity 28–47‰, temperature 10–30°C). — Perkins 1998: 95 (checklist of shallow-water polychaetes from Florida). — Bianchi & Morri 2001: 216 –218 (Orbetello Lagoon, Italy; reef-builders, competition with Hydroides dianthus (Verril, 1873) and ecological preferences; salinity 13–48‰, temperature 7–30°C). — Schwindt et al. 2001: 139 –147 (impacts on Mar Chiquita Lagoon, Argentina). — Schwindt et al. 2004: 111 –118 (physical effects on the same lagoon). — Bastida-Zavala 2008: 19, fig. 5A (California and Hawaii). — Carlton & Eldredge 2009: 60 –61 (Hawaii; invasion history and local economic impacts). — Ben-Eliahu & ten Hove 2011: 14 ( Israel, Suez Canal and Egypt; 0.3 m; under rocks, encrustation of tin can). — Cohen 2011: webpage (Los Angeles Harbor, California; small boat marina; four specimens). — Pernet et al. 2016: 13 –18 (Los Angeles River, California; established populations).

Material examined

143 specimens: CB (64) Sep. 2000, 2001 and Aug. 2012, CH (17) Sep. 2004, IR (1) Aug. 2005, TB (2) Jul. 2002, GB (12) Sep. 2002, SF (47) Sep. 2000, 2011 and 2012.

Additional material

25 specimens: LACMNH N8835 and N1873, many specimens (Lake Merritt, Oakland, California, 27 Jul. 1934 and 24 Jul. 1937); LACMNH N5141, 15 specimens (approx. 21°17' N, 157°50' W, Ala Wai Canal, Honolulu, Hawaii, 18 Apr. 1948, coll. R.W. Hiatt).

Diagnosis

This species is gregarious and can build large colonies. Tube white, often covered by a dark film of microalgae; with large peristomes (collar-like rings); without longitudinal ridges or alveoli ( Fig. 2E ). Opercular peduncle smooth, white. Operculum fig-shaped, with a brown, horny plate covered with black spines, curving inwards ( Fig. 2G ), sometimes with accessory spinules. Sometimes, the horny plate is completely covered by these spines. Thoracic membranes well developed, not fused dorsally ( Fig. 2F ). Special collar chaetae coarsely serrated.

Measurements: Total length = 11.1 mm (n = 14, range (r): 3.1–25.3, SD = 6.9); thorax length = 2.7 mm (n = 15, r: 1.4–3.8, SD = 0.9), thorax width = 0.9 mm (n = 15, r: 0.5–1.6, SD = 0.3); peduncle and operculum length = 1.7 mm (n = 15, r: 0.8–3.0, SD = 0.6); operculum length = 1.1 mm (n = 15, r: 0.6–1.6, SD = 0.3); operculum diameter = 0.7 mm (n = 15, r: 0.5–1.0, SD = 0.2).

Taxonomic remarks

Ficopomatus enigmaticus has a long historical record of invasions ( ten Hove & Weerdenburg 1978). Zibrowius (1992: 91) suggested that F. enigmaticus was probably introduced to Europe and San Francisco during the First World War (1914–1919), because the first records in Europe, including the original description ( Fauvel 1923), Rioja (1931) from Spain, and the specimens collected in 1931 from Lake Merrit, Oakland (revised by ten Hove & Weerdenburg 1978), were made a few years after the war. Zibrowius also rejected the hypothesis of Fauvel (1923) of an Indian-Indonesian origin. He thought it more likely that the species originates in a subtropical to temperate region, perhaps southern Australia. Given that five of the six species of Ficopomatus are present in Asia, this region seems a likely point of origin. Recently, Styan et al. (2017) analyzed several populations of Ficopomatus enigmaticus from southern Australia, using mitochondrial (Cyt B) sequencing and nuclear marker (iSSR) profiles, concluding that these populations represents cryptic sympatric species.

This species plays an important ecological role in some localities, because it builds large colonies in some Mediterranean ( ten Hove 1974; ten Hove & Weerdenburg 1978; ten Hove & van den Hurk 1993; Fornós et al. 1997; Bianchi & Morri 2001) and Patagonian coastal lagoons ( Schwindt et al. 2001, 2004), where it forms reefs or micro-atolls, 2–3m in diameter( ten Hove&van den Hurk1993; Schwindt et al. 2001, 2004), and can reach 750 m in length as in Lake Tunis ( ten Hove & van den Hurk 1993).

Recently, colonies( Fig.2H ) of this species were collected from Long Beach, California ( Pernet et al. 2016). Bastida-Zavala (2008) recorded F. enigmaticus from fouling plates in San Diego, California; however, this record was a mistake because the samples examined were actually from San Francisco, California.

Ecology

Intertidal to sublittoral (2 m). In lagoons with salinities of 13–48‰, temperature 7–30°C ( Bianchi & Morri 2001); on Poaceae grass, submerged wood, stones and shells, bivalves, bryozoans, rocky, sandy and artificial substrates such as tin cans ( Fauvel 1923; ten Hove & Weerdenburg 1978; Ben-Eliahu & ten Hove 2011) and settlement plates (this study).

Distribution

World-wide in temperate, brackish-water lagoons. Northern Europe, Mediterranean, Black Sea, Caspian Sea, South Africa, southern Australia, Japan, Hawaii, California, northern Gulf of Mexico, Uruguay and northern Argentina ( ten Hove & Weerdenburg 1978). In this work, Ficopomatus enigmaticus was found abundantly on fouling plates from Chesapeake Bay, Virginia (first recorded there in 1994 by L. McCann) and San Francisco Bay, California; and occasionally from Charleston, South Carolina, Indian River and Tampa Bay, Florida, and Galveston Bay, Texas ( Fig. 3 ). The material studied extends its eastward range from Rockport, Texas ( ten Hove & Weerdenburg 1978), to Pensacola Bay, Florida (750 km); our study also extends its southward range from Barnegat Bay, New Jersey ( Hoagland & Turner 1980) to Indian River, Florida (1500 km), though only one specimen was found there.

Figures

Fig. 2. Serpulids from United States fouling plates. Crucigera websteri. A. Operculum, juvenile from Humboldt Bay, California. B. Operculum, adult from San Pedro, California (LACMNH-N8819). – C. zygophora. C. Operculum, juvenile from Alaska (SERC). D. Operculum, adult from Canoe Bay, Alaska (LACMNH-N2128). – Ficopomatus enigmaticus. E. Tubes from Lake Merritt, California (LACMNH-N5141). F. Body, from Chesapeake Bay, Virginia (SERC-59327). G. Operculum, from Chesapeake Bay, Virginia (SERC-60530R). H. Colonies in Long Beach, California (photo by Bruno Pernet).– F. miamiensis. I. Tubes, from Chetumal Bay, Mexican Caribbean (ECOSUR). J–K. Operculum, from Galveston Bay, Texas (SERC-88344RF). – F. uschakovi. L. Tube, from Corpus Christi, Texas (SERC-88883). M. Thorax and operculum, from Galveston Bay, Texas (SERC-86995). N. Operculum, from La Encrucijada, Chiapas (UMAR-Poly 113). O. Thorax and operculum, from Corpus Christi, Texas (SERC-88883).

Fig. 3. Distribution of serpulids (Crucigera and Ficopomatus spp.) from United States fouling plates (closed symbols) and literature records (open symbols).