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Anthropogenic spread of the immigrant sea­spider Ammothea hilgendorfii (Arthropoda: Pycnogonida: Ammotheidae) in UK waters

Roger N. Bamber

Marine Biodiversity Records / Volume 5 / August 2012 / e78 DOI: 10.1017/S1755267212000668, Published online:

Link to this article: http://journals.cambridge.org/abstract_S1755267212000668

How to cite this article: Roger N. Bamber (2012). Anthropogenic spread of the immigrant sea­spider Ammothea hilgendorfii (Arthropoda: Pycnogonida: Ammotheidae) in UK waters. Marine Biodiversity Records,5, e78 doi:10.1017/S1755267212000668

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Downloaded from http://journals.cambridge.org/MBD, IP address: 86.169.136.60 on 31 Aug 2012 Marine Biodiversity Records, page 1 of 5. # Marine Biological Association of the United Kingdom, 2012 doi:10.1017/S1755267212000668; Vol. 5; e78; 2012 Published online Anthropogenic spread of the immigrant sea-spider Ammothea hilgendorfii (Arthropoda: Pycnogonida: Ammotheidae) in UK waters roger n. bamber ARTOO Marine Biology Consultants, Ocean Quay Marina, Belvidere Road, SO14 5QY, UK

The ammotheid pycnogonid Ammothea hilgendorfi was first recorded outside its native range of the Pacific Ocean in 1978 in southern and in the Lagoon of Venice in 1979 to 1981. Both occurrences are attributed to anthropogenic dispersion by shipping. Subsequently, it has spread further in the UK to and, most recently, to Essex. The world distribution of this species and the history of its occurrence outside the Pacific are reviewed, and hypotheses for its recent spread in the UK, via recreational boating, are presented.

Keywords: pycnogonids, Ammothea, anthropogenic introduction, alien, marinas

Submitted 20 May 2012; accepted 24 June 2012

INTRODUCTION DISTRIBUTION

The pycnogonid Ammothea hilgendorfi (Bo¨hm, 1879) Western Pacific records (Figure 1) is a common littoral to infralittoral ammotheid species around the of Japan, and has also long been The type locality of Ammothea hilgendorfi is at , known from the Pacific coasts of California and Mexico (for Honshu¯, Japan, at 5 to 7 m depth (Bo¨hm, 1879, as Corniger many years under the junior synonym Lecythorhynchus mar- Hilgendorfi gen. nov. sp. nov.), since when it has been recorded ginatus Cole, 1904). throughout Japan in littoral to shallow waters (Helfer, While normally found free-living in epizoan and epiphytic 1938, as Lecythorhynchus marginatus; Hedgpeth, 1949, as communities and in heterogeneous sediments, this species Lecythorhynchus; Utinomi, 1951, as Lecythorhynchus; Utinomi, has been recorded ectocommensally on the holothurian 1954, as Lecythorhynchus; Stock, 1954, as L. marginatus; Holothuria (Selenkothuria) moebii (Ludwig) (Ohshima, Utinomi, 1959, as Lecythorhynchus; Utinomi, 1971, as 1927, as Lecythorhynchus) and on the starfish Coscinasterias Lecythorhynchus; Nakamura & Child, 1983; Nakamura, 1987 acutispina (Stimpson) (Nakamura & Fujita, 2004). It is often [with synonymy and literature]; Nakamura & Child, 1991; associated with hydroids, such as species of Abietinaria Nakamura, 1994; Turpaeva, 2001, as Leionymphon marginatus; Kirchenpauer (e.g. Hedgpeth, 1940); the protonymphon Nakamura & Fujita, 2004). In addition, sparse records exist larvae have been found endoparasitically in association with from Korea (Kim, 1984; Kim & Hong, 1986; Hong & Kim, the campanulariid Orthopyxis everta (Clark) (Russel & 1987), from China (Lou, 1936, as Lecythorhynchus)andfrom Hedgpeth, 1990, as Eucopella everta). The overall recorded the east of Russia (Losina-Losinsky, 1933, as depth-range for A. hilgendorfi is 0 to 15 m. Lecythorhynchus marginatus), again in littoral to shallow waters. This Pacific species was discovered in the Lagoon of Venice in 1979 to 1981, where its presence was attributed to anthro- Mid-Pacific records pogenic introduction via ships’-hull fouling (Krapp & Sconfietti, 1983). In August 1978, two specimens were col- From mid-oceanic islands in the Pacific, Ammothea hilgen- lected in , UK (Bamber, 1985), on the south dorfi has been recorded twice in the Society Islands (Child, coast of England, the second record outside the Pacific; this 1970; Mu¨ller, 1989) as well as in Hawaii (Hilton, 1942, as occurrence of A. hilgendorfi was also attributed to anthropo- Lecythorhynchus ovatus sp. nov.). genic dispersion by shipping. Subsequent records in the UK, including its most recent discovery on the east coast, are dis- cussed herein. Eastern Pacific records Ammothea hilgendorfi has been recorded frequently along the coasts of California and Mexico (Cole, 1904, as

Corresponding author: Lecythorhynchus marginatus sp. nov.; Hall, 1913, as R.N. Bamber L. marginatus; Schmitt, 1934, as L. marginatus; Hedgpeth, Email: [email protected] 1940, as L. marginatus; Hedgpeth, 1941, in key, as

1 2 roger n. bamber

Ammothea hilgendorfi was subsequently found in large numbers in , Dorset (50841.8′N01859.0′W) 1 (mistakenly cited as Portland Harbour in Bamber, 2010). The initial record was in July 2001, amongst epifaunal fouling material on a marina pontoon; by 2006, A. hilgendorfi was regularly found under boulders around the Harbour, in densities ranging from ‘occasional’ to ‘super- abundant’. In Poole Harbour, it now (2011) seems to be present intertidally (mid- level) virtually wherever there is suitable habitat, i.e. under small, rough boulders lying on medium to coarse sand, where several tens of animals may aggregate (L. Baldock, personal communication). By 2009, it had also been recorded outside Poole Harbour, at South Beach, Studland Bay, where it was certainly absent in 2006. Meanwhile, by 2010, the population was well-established, with numerous intertidal records Fig. 1. Ammothea hilgendorfi (Bo¨hm, 1879), adult female (after Bamber, 2010). Dark stippling on the trunk and legs indicates dark-brown around the northern half of the particularly under pigmentation patches. boulders, but also subtidal records down the western shore to the mouth of the ria. In the same year Ammothea hilgen- dorfi was recorded elsewhere in the Solent, including L. marginatus; Hilton, 1943, as L. marginatus; Hedgpeth, Yarmouth on the (50843.0′N01835.5′W), 1951, as L. marginatus; Hedgpeth, 1975; Child, 1979; again amongst epifaunal fouling on a marina pontoon. In Hedgpeth & Haderlie, 1980; Russel & Hedgpeth, 1990; Bain, the spring of 2011, large numbers of A. hilgendorfi were 1991). taken from pontoon epifauna in Ocean Quay Marina, the site of the old Southampton Outer Dock (50853.7′N 01823.3′W), as well as elsewhere in Southampton Water (Horton et al., 2011). OCCURRENCE AS A XENOBIONT In 2010, specimens of Ammothea hilgendorfi were collected for the first time from the southern North Sea on the east coast The first published record of Ammothea hilgendorfi outside of England. During a benthic survey of the River Blackwater the Pacific, as an anthropogenically-introduced species, was in Essex, in the vicinity of Bradwell Power Station by Krapp & Sconfietti (1983; see also Candela et al., 1983), (51844.7′N0853.5′E), A. hilgendorfi was the second- who discovered A. hilgendorfi in the Lagoon of Venice most-common of nine species of pycnogonid collected, with amongst fouling communities in 1979 to 1981, in nearly fully- twenty individuals taken at six sites. These specimens were marine conditions. These authors attributed its presence to collected in grab samples from sedimentary habitats compris- arrival from East Asian waters in the hull-fouling of ships: ing mud with fine sand and shell, at depths between 6 and there is an extensive shipping trade from the Far or Middle 31.5 m. The Blackwater Estuary is also an area of recreational East via the Suez Canal to the port of Venice (e.g. Galil, yachting, with numerous moorings around Mersea Island to 2000; Streftaris et al., 2005). There have been no subsequent the north of the sampling area and at Bradwell Marina to records of A. hilgendorfi on the adjacent shores of the the south-east. This area was subject to some intensive survey- Adriatic Sea. ing in relation to existing and proposed power stations from United Kingdom records are listed in Table 1, with a map the 1960s to the mid-1980s (e.g. Bamber & Henderson, showing the key locations as Figure 2. 1981), and again in the late 1990s: A. hilgendorfi was certainly Ammothea hilgendorfi was first recorded in UK waters by absent then. Bamber (1985), from two immature specimens collected in August 1978 in Southampton Water, Hampshire (50849.5′N01819.3′W), in fully marine conditions. Southampton Water is a ria on the south coast of England, DISCUSSION about 13 km long and 2 km wide at its widest; its southern (seaward) end connects with the Solent, a separating Pycnogonids do not have an obligate dispersive phase in their the Isle of Wight from the Hampshire mainland, while at its life history. A few species have been observed swimming (e.g. northern end lies Southampton, a major port with an exten- Cole, 1901; Fage, 1932; Clark & Carpenter, 1977), but this sive shipping trade, including with Japan. This area of coast- appears to be a method of maintaining a position above the line now supports resident populations of a number of sea-bottom rather than for directional movement; it may Japanese species, including the tunicate Styela clava allow for tidal dispersion (Morgan, 1977, 1978). Thus, achiev- Hedman and the serpulid tube-worms Pileolaria (P.) berke- ing a wide distribution requires either a long time, or some leyana (Rioja) and Hydroides ezoensis Okuda, although the assistance in dispersal, either in drifting algae or by anthropo- latter may have crossed the from French genic transport. Pycnogonids have been recorded in coasts where the Japanese oyster Crassostrea gigas ships’-hull fouling (e.g. Bamber, 1979), a habitat in which (Thunberg) had been transplanted. The establishment of they might be expected to be able to complete their life-cycle. A. hilgendorfi in Southampton Water was first confirmed with the discovery of an adult in October 1987 (Bamber, 1Searches up until the end of 2010 have failed to find A. hilgendorfi in 1988). Portland Harbour (L. Baldock, personal communication). spread of ammothea hilgendorfi in uk waters 3

Table 1. Dates and locations of Ammothea hilgendorfi records in UK waters.

Area Year Location Latitude–longitude

Southampton Water, Hampshire 1978 Fawley Marsh 50849.51′N 001819.37′W 1987 Fawley Marsh 50849.51′N 001819.37′W 2010 Marchwood 50844.07′N 001826.38′W 2010 Victoria Country Park 50852.20′N 001821.10′W 2010 Estuary 50853.28′N 001824.33′W 2010 Western Shore 50852.94′N 001822.05′W 2010 Western Shelf 50853.38′N 001823.01′W 2010 Hamble Point 50850.99′N 001818.94′W 2011 Ocean Village Marina 50853.78′N 001823.35′W West Solent 2010 Yarmouth Marina 50842.30′N 001830.19′W 2010 Pennington to Lepe 50844′N 001828′W 2010 Stanswood Bay 50848′N 001818′W Poole Harbour, Dorset 2001 Poole Quay 50842.86′N 001859.61′W 2006 Baiter 50842.61′N 001858.51′W 2006 Poole Ferry 50842.71′N 001858.06′W 2006 Poole Park 50842.79′N 001857.95′W 2006 Pierce Avenue 50842.58′N 001857.43′W 2006 Lilliput 50842.06′N 001856.69′W 2006 Brownsea Island 50841.22′N 001859.05′W Poole Bay, Dorset 2009 South Beach, Studland 50838.49′N 001856.37′W Blackwater Estuary, Essex 2010 Bradwell 51844.73′N 000853.46′W

As well as being the location of a major port, Southampton commercial and tourism activities, and cites lagoons, Water and the adjacent Solent house the greatest concentration and marinas as habitats particularly susceptible to invasions. of marinas and recreational yachts in the UK. Poole Harbour is It thus seems likely that A. hilgendorfi was first introduced also a significant site for yacht marinas, and in easy sailing dis- to the UK by international shipping into Southampton, and tance from Southampton. It is notable that the majority of the has subsequently spread westwards along the south coast to records of A. hilgendorfi from sites in Dorset, Hampshire and Dorset and eastwards to Essex with the assistance of rec- the Isle of Wight are from marinas. The River Blackwater reational yachting. Other likely sites for the appearance of Estuary is home to some commercial fishing (trawling) and a this species are Brighton (East Sussex) and the Channel nuclear power station, but currently is largely a centre for Islands, both common destinations for short-distance rec- recreation, notably with yacht marinas. Galil (2000) notes reational yachting from the Solent area. that the increase in spread of xenobiont species in the Climatically, the UK is amenable to a species originating in Mediterranean is associated with increases in seafaring, Japan, as can be seen from the current distribution of the

Fig. 2. Map of southern England showing locations of the UK records of Ammothea hilgendorfi. 4 roger n. bamber

tunicate Styela clava around the British Isles (Davis & Davis, Clark W.C. and Carpenter A. (1977) Swimming behaviour in a pycnogo- 2004). It appears to have taken A. hilgendorfi some two nid. New Zealand Journal of Marine and Freshwater Research 11, decades to spread from Southampton to Poole Harbour 613–615. (where it has since proliferated), and three decades to reach Cole L.J. (1901) Notes on the habits of pycnogonids. Biological Bulletin. the southern North Sea coast of England; the main dispersion Marine Biological Laboratory, Woods Hole 2, 195–207. (so far) of this species in the UK seems to have occurred Cole L.J. (1904) Pycnogonida of the west coast of North America. during the past decade. It will be interesting to see how long Harriman Alaska Expedition 10, 249–330. pls XI–XXVI. it takes to achieve a similar spread around the UK as has Styela clava. Fortunately, while a successful Davis M.H. and Davis M.E. 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