The Australian Distribution of the Introduced Sphaeromatid Isopod, Paracerceis Sculpta

THE AUSTRALIAN DISTRIBUTION OF THE INTRODUCED SPHAEROMATID ISOPOD, PARACERCEIS SCULPTA BY CHAD L. HEWITT 1,2 / and MARNIE L. CAMPBELL 1,3 / 1 / CSIRO Marine Research, Centre for Research on Introduced Marine Pests, GPO Box 1538, Hobart, Tasmania 7001, Australia ABSTRACT The sphaeromatid isopod, Paracerceis sculpta, identiŽed for the Žrst time in Australia at Townsville, Queensland, in 1982 (from a collection made in 1975) is a native of the PaciŽc Coast of North and Central America and has a wide global introduced distribution. We report on the distributionof P. sculpta in Australia based primarily on a series of port surveys speciŽcally designed to detect introduced fauna and supplemented by museum records. Based on its current distribution and the local abundance at speciŽc sites, P. sculpta is primarily associated with and likely to be transported by hull fouling, although it may rarely be transported by ballast water. Paracerceis sculpta is likely to continue its expansion in Australian waters either through coastal (domestic) translocations or directly from overseas. RÉSUMÉ L’Isopode Sphaeromatidae, Paracerceis sculpta, identiŽé pour la première fois en Australie à Townsville, Queensland, en 1982 (à partir d’une récolte faite en 1975) est originaire de la côte PaciŽque de l’Amérique du Nord et Centrale et présente une large répartition globale comme espèce introduite. Nous nous intéressons ici à la répartition de P. sculpta en Australie, à partir d’une série de contrôles portuaires spéciŽquement destinés à la détection de la faune introduite, et complétés par des références de collections muséologiques. À partir de sa répartition habituelle et de son abondance locale sur des sites spéciŽques, P. sculpta est principalement associé à et probablement transporté par les coques de navires, bien qu’il puisse, de temps en temps, être transporté par les eaux de ballast. Paracerceis sculpta va probablement poursuivre son expansion dans les eaux australiennes, soit par les transports côtiers (domestiques), soit directement de l’étranger. 2 / Fax: +61.362325487; e-mail: Chad.Hewitt@marine.csiro.au 3 / Current address: Environment Australia, Marine Protected Areas Section, Canberra, ACT 2601, Australia. c Koninklijke Brill NV, Leiden, 2001 ° Crustaceana 74 (9): 925-936 926 CHAD L. HEWITT & MARNIE L. CAMPBELL INTRODUCTION The isopod, Paracerceis sculpta (Holmes, 1904) is native to the Northeast PaciŽc from southern California (Miller, 1968) to Mexico (Brusca, 1980; Menzies, 1962). Paracerceis sculpta is associated with algae (Pires, 1980) and calcareous sponges, often living in the choanocyte chambers (Shuster, 1987). These associations with common benthic fouling organisms (algae and sponges) may be the main reason why the species has been implicated in translocation via hull fouling (Harrison & Holdich, 1982), although Poore & Storey (1999) also consider the potential for transport via ballast water as high. Paracerceis sculpta has been introduced to Brazil (Loyola e Silva et al., 1999; Pires, 1980, 1981), Hawaii (Coles et al:; 1997; 1999I Miller, 1968), the Atlantic coast of Europe (Spain: Rodriguez et al., 1992), the Mediterranean (Tunisia: Rezig, 1978; Italy: Forniz & Maggiore, 1985; Forniz & SconŽetti, 1985), Hong Kong (Bruce, 1990), and Australia (Harrison & Holdich, 1982; Hass & Jones, 1999; Hass & Knott, 2000) (Žg. 1). Paracerceis sculpta specimens collected from a pontoon in the Townsville port area in 1975 are the Žrst record of this species in Australian waters (Harrison & Holdich, 1982). The authors suggested trans-PaciŽc shipping as the vector responsible for the species’ introduction to Townsville. This supposition is now further supported by recent collections of P. sculpta in other international Žrstports-of-call in temperate Australia. It has been reported as widespread in Western Australia, particularly Bunbury, Mandurah (Peel Inlet), and Fremantle/Swan River (Hass & Knott, 2000), and Port Denison (Hass & Jones, 1999) (Žg. 2). Fig. 1. Published world-wide distribution of Paracerceis sculpta (Holmes, 1904). Native range is denoted by and introduced locations are denoted by . AUSTRALIAN DISTRIBUTION OF PARACERCEIS SCULPTA 927 Fig. 2. Australian distribution of Paracerceis sculpta (Holmes, 1904) based on port surveys or museum collections.Symbols represent National Port Survey localities where P. sculpta was detected at high densities ( ), low densities (<10 individuals collected in the port) or limited distribution( ), or absent ( ); P. sculpta reports from the literature or other communications denoted by . This study records the distribution of Paracerceis sculpta in several new localities based upon the Žndings of a series of port baseline surveys in Australia and discusses the likely introduction vectors based on analyses of local distribution and abundance. MATERIALS AND METHODS In 1995, a National Port Survey Program to detect introduced species was established by CSIRO Marine Research, Centre for Research on Introduced Marine Pests (CRIMP) in collaboration with the Australian Association of Ports and Marine Authorities. This continental-wide Program is designed to establish a series of baseline evaluations for introduced species targeting prioritized sample locations such as active and disused berths, marinas and Žshing vessel moorings, and shipping channel markers using a standardized suite of sampling protocols detailed in Hewitt & Martin (1996, 2001). In brief, all biological material from hard substrates is sampled by removing all biomass in 0.10 m2 quadrats (size selected based on previous studies for encrusting communities, see Hewitt & Martin, 1996, 2001) at depths of ¡0.5, ¡3, and ¡7 m Mean Low Water (MLW). Quadrat samples are replicated three times per depth at all sites (with the exception of channel markers). In addition, qualitative samples are collected from rocky ‘rip-rap’, breakwalls, bridge abutments, and other 928 CHAD L. HEWITT & MARNIE L. CAMPBELL artiŽcial substrates both inside and adjacent to port areas. The independence of occurrence (presence at a site) and prevalence (percentage occurrence in samples at a site) of Paracerceis sculpta at hull fouling versus ballast water related sites was analysed with R £ C tests of independence (Sokal & Rohlf, 1981). G statistics were adjusted due to small sample sizes. To date, CRIMP has undertaken 16 surveys independently (Mackay and Hay Point, Queensland (QLD); Williamstown/Melbourne and Geelong, Victoria (VIC); Lady Barron and Devonport, Tasmania (TAS); and Bunbury and Port Hedland, Western Australia (WA)) or in collaboration with universities and state/territorybased organizations (Darwin, Northern Territory (NT) with the Museum and Art Gallery of the Northern Territory; Gladstone, QLD with the Central Queensland University; Newcastle, Botany Bay, Port Kembla (and Wollongong) and Eden, New South Wales (NSW) with New South Wales Fisheries Research Institute; Port Lincoln, South Australia (SA) with Flinders University; and Fremantle, WA with Murdoch University). Following the detection of Paracerceis sculpta in the port surveys, identiŽcations were conŽrmed by G. C. B. Poore (Museum Victoria), specimens were lodged with the Tasmanian Museum (TMH) and queries were made of Australian State and Territory museum collections to determine if additional specimens had been collected and identiŽed, but not reported. Museum reference numbers are provided for all lots identiŽed from Australia. RESULTS Baseline Port Surveys Paracerceis sculpta was not detected in Mackay, QLD; Newcastle, NSW; Geelong, VIC; Devonport and Lady Barron, TAS; Port Lincoln, SA; Port Hedland, WA; or Darwin, NT (Žg. 2). A brief description of the P. sculpta populations that were found at each of the National Port Survey localities is provided below. Tropical Australian ports. — Hay Point, QLD (21± 150 S 149± 180 E): one ®-male was collected from a single pile scraping (from a total of 6 quadrats) at ¡3 m MLW in the tug harbour at the Port of Hay Point on 17 May 1997 (Museum Ref. No.: TMH G4092). No additional specimens were collected from either the Hay Point or adjacent Dalrymple Coal Terminals (57 quadrats at six sites). The area experiences a high silt load and water depths range from 0 to ¡5 m MLW. Samples collected from the nearby Port of Mackay (<20 km north) did not contain P. sculpta (Žg. 2). Gladstone, QLD (23± 510 S 151± 190 E): one ®-male was collected from a pile scraping (¡7 m MLW) at South Trees Wharf on 17 March 2000 (Museum Ref. AUSTRALIAN DISTRIBUTION OF PARACERCEIS SCULPTA 929 No.: TMH G4113). No additional specimens were collected out of 8 other quadrats at this site, or at any of the 87 quadrats from 17 other sample sites in the port. Temperate Australian ports. — Williamstown, Port Phillip Bay, VIC (37± 490 S 145± 400 E): two ®-males and 1 female were collected from a pile scraping at Anne Street Pier, Williamstown, <10 m MLW, on 19 September 1995 (Campbell & Hewitt, 1999) (Museum Ref. No.: TMH G4114: 1 ®-male; and MoV J41569: 1 ®-male and 1 female). This pier is the property of the Australian Navy. Eden, NSW (37± 050 S 149± 540 E): eight ®-males were collected from pile scrapings at ¡3 and ¡7 m MLW depths, on Wharf C (the Quarantine Wharf for international vessels) on 6 November 1996 (Museum Ref. No.: TMH G4111: 5 ®-males). No other specimens were recorded from 39 quadrat samples at six additional sites. Paracerceis sculpta was not detected in 1984-85 during a previous survey of Eden by Hutchings et al. (1989). That survey did not sample in the wharf area of Eden where these specimens were collected. Botany Bay, NSW (34± 000 S 151± 140 E): Paracerceis sculpta was collected in Botany Bay between 19 and 21 October 1998 during a baseline port survey for introduced species (Pollard & Pethebridge, 2000). A single ®-male of Paracerceis was found at a one site, Lady Robinsons Beach, on rocky ‘rip-rap’, at <2.5 m MLW depth. No other specimens were recorded from either 96 quadrat samples or 10 qualitative collections at 24 additional sites based on identiŽcations by G. Poore (Museum of Victoria). Wollongong Harbour, NSW (34± 250 S 150± 540 E): Paracerceis sculpta was collected in Wollongong Harbour on 17 May 2000 during the baseline port survey of Port Kembla (G. C. B. Poore, pers. comm.). A single ®-male and two juvenile specimens of Paracerceis were found in Belmore Basin, a Žshing vessel berthing area, at 0.5 m MLW. No other specimens were recorded from either seven other quadrat samples or qualitative collections in the harbour, or in 116 quadrat samples and nine qualitative samples at 33 additional sites in Port Kembla Harbour, 6 km to the south. Fremantle, WA (32± 020 S 115± 440 E): Paracerceis sculpta is widespread in the Swan River, the Fremantle small boat marinas and in the port areas of Cockburn Sound (Žg. 3). Collections were made from 18 April to 14 May 1999 during a baseline evaluation of the port (Museum Ref. No.: TMH G4115: 3 ®-males and 1 female). Paracerceis was found at 10 localities and was collected from pile scrapings at depths varying from ¡0.5 to ¡7 m MLW. With the exception of the Cockburn Sound sites, all locations where Paracerceis occurred were associated with small vessel usage (i.e., Žshing vessels and recreational boats) (Žg. 3). Bunbury, WA (33± 190 S 115± 390 E): Paracerceis sculpta is widespread in the Bunbury port region (inner and outer harbours) and in the Koombana Bay shipping channel at the entrance to the port (Žg. 4). Collections were made from 2 to 930 CHAD L. HEWITT & MARNIE L. CAMPBELL Fig. 3. Distribution of Paracerceis sculpta (Holmes, 1904) in The Port of Fremantle and Cockburn Sound, WA. Symbols represent sample sites. Symbol sizes are proportionalto prevalence (percentage of samples at a site, see legend) of P. sculpta; open circles represent absence ( ); Žlled circles represent locations where P. sculpta was detected at: commercial or naval berths ( ); and recreational or Žshing vessel berths and moorings ( ). See Žg. 2 for location of Fremantle in Australia. 6 March 1996 (Museum Ref. No.: TMH G 4112: 8 ®-males). Paracerceis was found at eight localities at depths varying from ¡0.5 to ¡7 m MLW. The majority of individuals were collected from pile scrapings, although some were taken in qualitative samples associated with sponges in the rocky ‘rip-rap’. Maximum densities were estimated at 720 individual females m¡2 (inner harbour) and 160 individual males m¡2 (outer harbour). In the two high-abundance and wide-distribution ports (Fremantle and Bunbury, WA), no signiŽcant difference in Paracerceis sculpta occurrence at commercial berths (34.6%) and recreational and Žshing vessel berths (39.1%) was detected (Gadj 1 D 0:103, P > 0:05). In contrast, the prevalence (total occurrence in samples from these sites) was found to be signiŽcant (Gadj 1 D 12:06, P < 0:01). Paracerceis was found in 25.3% of the samples from commercial berths and AUSTRALIAN DISTRIBUTION OF PARACERCEIS SCULPTA 931 Fig. 4. Distribution of Paracerceis sculpta (Holmes, 1904) in The Port of Bunbury, WA. Symbols represent sample sites with size proportional to Paracerceis prevalence (percentage of samples at a site, see legend); open circles represent absence ( ); Žlled circles represent locations where P. sculpta was detected at: commercial or naval berths ( ); and recreational or Žshing vessel berths and moorings ( ). See Žg. 2 for location of Bunbury in Australia. channel markers, as opposed to 55.6% of samples from recreational and Žshing vessel berths, slipways, and tug basins. Museum collections A search was made of all Australian State and Territory Museum collections for specimens of Paracerceis sculpta: Queensland Museum (QM). — Hayle’s Wharf, Ross Creek (incorrectly identiŽed as Ross River Creek on the label), Townsville, QLD (19± 160 S 146± 490 E): a single lot of Paracerceis sculpta (Museum Ref. No.: QM W7968: 1 ®-male and 1 non-ovigerous female) collected by P. Otteson, October 1975, from ‘the side of a pontoon’. These specimens are the basis of the identiŽcation of P. sculpta in Australia by Harrison & Holdich (1982). Hayle’s Wharf has always been the ferry terminal between Townsville and Magnetic Island. Museum and Art Gallery of the Northern Territory (MAGNT). — None (G. Daley, pers. comm.). Western Australian Museum (WAM). — One lot (2 ®-males and 2 females) (Museum Ref. No.: WAM C.23303), collected by C. Hass from a jetty pile at <0.5 m MLW depth amongst fanworms and ascidians, in Bunbury Harbour, 1 October 1997. South Australian Museum (SAM). — One lot, New Haven Marina, Adelaide, SA (34± 450 S 138± 300 E) (Museum Ref. No.: SAM C 5930: 2 ®-males and 3 nonovigerous females), collected by T. Laperousaz, May 2000, from the side of a pontoon of a Žsherman’s wharf at 0.5 m depth among algae. 932 CHAD L. HEWITT & MARNIE L. CAMPBELL No additional lots have been identiŽed from the Museum’s collections (T. Laperousaz, pers. comm.). Museum Victoria (MoV). — One lot (1 ®-male and 1 female) from the CRIMP Port Phillip Bay survey, Anne Street Pier, Williamstown (mislabeled as from Geelong) (Museum Ref. No.: MoV J41569). No additional lots were identiŽed from the Museum’s collections prior to 1998 (Poore & Storey, 1999). Tasmanian Museum (TMH). — Several lots from the CRIMP survey collections (see above) have been registered with the Tasmanian Museum but are maintained in that collection. Collection maintained and housed with CSIRO Marine Research in Hobart, Tasmania. Australian Museum (AM). — One lot, Store Beach, North Harbour, Port Jackson, NSW (33± 48:80 S 151± 17:20 E), (2 ®-males and 2 females; determined N. L. Bruce, Museum Ref. No.: AM P 54249), (P. Berents, pers. comm.). These specimens were collected 26 September 1984 by R. Springthorpe and J. Just at ¡9 m MLW depth, found in sponge on algae. It is unknown whether these specimens represent a larger population in Port Jackson. (N.B.: Additional specimens have been collected from Manley and Blackwattle Bay as part of a settlement plate study in March 2001 by CLH.) DISCUSSION Paracerceis sculpta is now known from thirteen localities in both tropical and temperate Australia. The species is associated with fouling communities in Australian harbours that have both low (e.g., Koombana Bay channel marker in Bunbury, and Cockburn Sound, WA; Eden and Botany Bay NSW) and high silt loads (e.g., Hay Point Tug Harbour, and Gladstone, QLD; Wollongong Harbour, NSW). The presumed native range of P. sculpta extends from tropical to subtropical East PaciŽc waters with temperatures ranging from 9± to 33± C. It survives as an introduced species in regions that are both tropical (e.g., Hawaii; Hay Point and Gladstone, QLD, Australia) and temperate (southern Brazil; Adriatic Sea; Williamstown, Port Phillip Bay, VIC; Botany Bay and Eden, NSW; Fremantle and Bunbury, WA). This wide tolerance to water temperature and silt load are likely to be important factors that enable it to survive cross equatorial transit and then survive in disturbed, frequently turbid, habitats such as those that occur in port environments. In the majority of sites (85%) where Paracerceis sculpta has been detected in Australia, it has been in low numbers (<10 detected from multiple sampling sites within a port) or with limited distribution (Townsville, Hay Point and Gladstone, QLD; Port Jackson, Botany Bay, Wollongong Harbour and Eden, NSW; AUSTRALIAN DISTRIBUTION OF PARACERCEIS SCULPTA 933 Williamstown, VIC; North Arm Marina, SA; Peel Inlet and Port Denison, WA). The breeding status of these populations is unknown. Of these sites, only Port Jackson, NSW and Williamstown, VIC, have both ®-males and females, while the remainder only have ®-males. This suggests that these populations represent either: (i) marginal populations; (ii) recent invasion events; or (iii) failing invasion events. The hypothesis that these are marginal populations is not supported by the regional distribution of sites either in Australia or of other parts of the world. It is not possible to speculate on the relative merits of the remaining hypotheses without temporal distribution data. Regardless, the likelihood of P. sculpta spreading to other harbours and marinas appears high. No single translocation mechanism is likely to be responsible for all Australian inoculations. Hull fouling has been implicated as the most likely vector for Paracerceis sculpta by numerous authors (Harrison & Holdich, 1982; Forniz & Maggiore, 1985; Rodriguez et al., 1992; Hass & Jones, 1999; Hass & Knott, 2000) and Hass & Jones (1999) collected material from recreational and Žshing vessel hulls. The hypothesis that P. sculpta is transported by hull-fouling communities is supported by our indirect evidence based on distributions and abundances in well-surveyed ports. Eight of the 11 sites in which P. sculpta was detected in low numbers or limited distribution, were primarily associated with recreational vessels (Port Jackson, NSW; New Haven Marina, SA; and Peel Inlet, WA), Žshing vessels (Wollongong Harbour and Eden, NSW; and Port Denison, WA) or slow moving vessels such as tugs (Hay Point Tug Harbour, QLD), ferries (Hayles Wharf, Townsville, QLD) and barges. These classes of vessels often have long residence times in port resulting in heavy fouling. This supports the hypothesis that hull fouling is the likely introduction vector in these locations. In order for a species to be transported by ballast water either adults or juveniles must be entrained from the plankton. Paracerceis sculpta is a sedentary species occupying the internal cavities of sponges (Shuster, 1987, 1992a, b) with no planktonic phase in its life history. It is conceivable that adults swimming adjacent to berth piles, or swept off as tychoplankton (benthic organisms swept into the water column by currents or other dislodging actions) could be drawn into the ballast tanks. Regardless, Williams et al. (1988) and Carlton & Geller (1993) document numerous benthic isopods and amphipods in ballast tanks. Therefore, while ballast water is an unlikely vector for transport of P. sculpta, two out of the 11 sites with either low numbers or limited distribution of P. sculpta receive discharge of ballast water. These two sites, where single specimens were collected, are the active commercial shipping berth at South Trees Wharf in Gladstone, QLD and the naval pier (Anne Street Pier) of Williamstown, VIC. 934 CHAD L. HEWITT & MARNIE L. CAMPBELL In Fremantle and Bunbury, WA, where Paracerceis sculpta was collected in high densities in a number of locations, no signiŽcant differences between the percent occurrence at recreational/Žshing vessel areas (hull fouling) and commercial (ballast water) were detected. Fifty percent of the sites where P. sculpta was collected in these two ports were potentially subjected to ballast discharge from commercial or naval shipping. In contrast, 63% of all sites where P. sculpta was collected were associated with Žshing/recreational vessels or slow moving vessels (note that at some of the commercial shipping sites there is overlap with Žshing/recreational/slow moving vessel activity). Paracerceis is signiŽcantly more prevalent (percentage occurrence in samples) at hull fouling associated sites than at sites with ballast discharge. With the exception of a single commercial berth in Inner Bunbury Harbour and the channel marker immediately outside the Inner Bunbury Harbour entrance (Žg. 4), the incidence of P. sculpta in commercial areas is low (present in 25% of quadrat samples from invaded commercial berths). In contrast, Paracerceis was detected in 56% of quadrat samples in Fremantle boat harbours, and at the Bunbury slipway (small vessels <500 metric tonnes) and disused jetty sites (commonly used for Žshing vessel mooring) suggests that small vessel hull fouling may have contributed to its local (within a port) and regional (Western Australia) distribution. Currently there is no indication that Paracerceis sculpta establishes populations outside of port and harbour environments. This may be a function of search effort, a limited dispersal capability of P. sculpta, or alternately, evidence of susceptibility of port environments to invasions. The surveys described herein negate search effort as a causal factor. Peracarid isopods, by nature, have limited dispersal capability, however, recreational and Žshing vessels frequently travel to non-port environments suggesting that inoculation of P. sculpta to these locations is likely. The potential for P. sculpta to be spread both by hull fouling and ballast water, and its capacity to survive and establish in diverse port environments, suggests that this species will continue to expand its distribution in Australian waters. ACKNOWLEDGEMENTS We would like to acknowledge the keen eye of Mr. Alec Moore for his initial identiŽcation of Paracerceis amidst numerous specimens. We are grateful for helpful information and aid from Christina Hass, Karen Gowlett-Holmes, and various museum workers including Peter Davie (QM), Gavin Daley (MAGNT), Diana Jones (WAM), Thierry Laperousaz (SAM), Gary Poore (MoV), and Penny Berents (AM). 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