Managing an Invasive Corallimorph at Palmyra Atoll National Wildlife Refuge, Line Islands, Central Pacific
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Biol Invasions https://doi.org/10.1007/s10530-018-1696-1 ORIGINAL PAPER Managing an invasive corallimorph at Palmyra Atoll National Wildlife Refuge, Line Islands, Central Pacific Thierry M. Work . Greta S. Aeby . Benjamin P. Neal . Nichole N. Price . Eric Conklin . Amanda Pollock Received: 29 September 2017 / Accepted: 26 February 2018 Ó The Author(s) 2018. This article is an open access publication Abstract In 2007, a phase shift from corals to pulverized bleach may be an effective tool to eradicate corallimorpharians (CM) centered around a shipwreck CM on a local scale. If applied strategically, partic- was documented at Palmyra Atoll, Line Islands. ularly in heavily infested ([ 66% cover) areas, active Subsequent surveys revealed CM to be overgrowing intervention such as this could be an effective the reef benthos, including corals and coralline algae, management tool to reduce CM impact on localized potentially placing coral ecosystems in the atoll at risk. areas and decrease colonization rate of remaining This prompted the U.S. Fish and Wildlife Service, the reefs. This is the first documentation of the response of lead management agency of the atoll, to remove the an invasive cnidarian to shipwreck removal. While shipwreck. Subsequent surveys showed reductions in this was a singular event in Palmyra, the spatial and CM around the ship impact site. We explain patterns temporal patterns of this invasion and the eradications of spread of the CM in terms of both life history and lessons described herein, are useful for anticipating local currents and show with a pilot study that and controlling similar situations elsewhere. Keywords Corralimorpharian Á Invasive species Á T. M. Work (&) Shipwreck disturbance Á Invasion dynamics Á Coral US Geological Survey, National Wildlife Health Center, reefs Á Control Á Management Á Phase shifts Honolulu Field Station, PO Box 50187, Honolulu, HI 96850, USA e-mail: [email protected] G. S. Aeby Introduction Hawaii Institute of Marine Biology, 46-007 Lilipuna Rd, Kaneohe, HI 96744, USA Coral reefs are subject to damage from both natural B. P. Neal Á N. N. Price and anthropogenic causes, including large scale storm Bigelow Laboratory for Ocean Sciences, 60 Bigelow Dr, events, temperature anomalies and subsequent bleach- East Boothbay, ME 04544, USA ing events, disease outbreaks on foundational species, E. Conklin overfishing, and ship groundings. The resultant dis- The Nature Conservancy-Hawaii, 923 Nuuanu Ave, ruption from these events can facilitate colonization or Honolulu, HI 96817, USA expansion by invasive organisms that can result in phase shifts (Chadwick and Morrow 2011; Hughes A. Pollock US Fish and Wildlife Service, Refuges, et al. 2010). Phase shifts are defined here as ‘‘extensive PO Box 50167, Honolulu, HI 96850, USA decreases in coral cover coinciding with substantial 123 T. M. Work et al. increases in some alternative benthic organism, due to capable of year-round reproduction via sexual (pelagic a pulse or press disturbance, that have per- spawning) or asexual clonal replication (Chadwick- sisted [ 5 year’’ (Norstro¨m et al. 2009). For example, Furman and Spiegel 2000; Chen et al. 1995). Some mass mortality of the sea urchin, Diadema antillarum, species of CM can rapidly replicate with a polyp throughout the Caribbean from an apparent disease doubling time in as little as 2 months (Chadwick and outbreak contributed to a phase-shift from coral- to Adams 1991). R. howesii is considered native to algal-dominated communities (Lessios 2016). In the Palmyra based on observations of its presence around Indo-Pacific, similar phase-shifts have occurred in the atoll prior to the outbreak (Work et al. 2008). response to shipwrecks where coral dominated com- Perhaps life history characteristics of the CM popu- munities change to those dominated by opportunist lation around the shipwreck contributed to the benthic organisms. On Rose Atoll in American Samoa observed rapid spread on the atoll. The increased where a fishing vessel wrecked on the reef, the percent presence of CM around the ship would be analogous to cover of turf/cyanobacteria was an order of magnitude an outbreaks of other native species such as the crown higher (40%) in areas surrounding the wreck as of thorns seastars on coral reefs (Pratchett et al. 2014). compared with reference sites on the same island In 2013, the FWS secured funding to disassemble (Schroeder et al. 2008). In the Line Islands, iron and remove the shipwreck at Palmyra Atoll NWR with enrichment from shipwrecks fueled growth of invasive all the metal shipped back to the mainland U.S. for cyanobacteria and turf algae (Kelly et al. 2012). Once scrap. Our objectives were to (1) document the established, invasive organisms are difficult to control, distribution of CM at Palmyra before and after with failures mostly due to late detection resulting in shipwreck removal, (2) examine the life history of sufficient time elapsing for invasives to occupy areas CM and the oceanographic conditions that could help too large for eradication to be practical (Hewitt and explain the observed patterns of spread, and (3) test Campbell 2007). However, in the rare attempts that methods for control CM. have been made to manage marine invasives, suc- cesses do exist; for instance the complete elimination of the invasive algae, Caulerpa racemosa near San Methods Diego, California. This effort was successful because the algae were detected early, were within a well- Surveys defined area, and were eradicated before they spread over a large scale (Anderson 2005). In 2011 and 2016, surveys of CM infestation were In 2007, a corallimorpharian (CM), Rhodactis conducted following protocols described previously howesii, was discovered smothering coral reefs at (Work et al. 2008). Briefly, a snorkeler was towed at Palmyra Atoll National Wildlife Refuge (Palmyra), a an approximate speed of ca. 50 m/min over benthic remote atoll within the Line Islands ca. 5° north of the transects that originated from the shipwreck site Equator (Work et al. 2008). The CM invasion towards the 8 major points of the compass (N, NE, encompassed a well-defined 1 km2 area with a E, SE, S, SW, W, NW). Every minute (ca. 50 m), the predominant NW to SE orientation surrounding a towboat stopped, and the snorkeler subjectively inte- longline vessel that had wrecked on the atoll in 1991. grated benthic cover during the tow scoring the level The distribution of the CM around the shipwreck and of CM infestation based on percent cover using the around mooring buoys fastened by iron chains following 0–3 scale: 0 = no CM; 1 = ca. 1–33%; suggested that substances leaching from metals, 2 = 34–66%; and 3 = [ 67% CM cover. Visual area possibly iron, might in part be driving the spread of covered by each 1 min tow was ca 500 m2.Tows the infestation on the reef (Work et al. 2008). In continued until at least three consecutive stops scored response, the US Fish and Wildlife Service (FWS) in at 0 were obtained or until it became too deep collaboration with The Nature Conservancy switched (20–25 m) to see the benthos from the surface or too out existing iron mooring chains with stainless steel in shallow (\ 1 m) for effective vessel operation. To 2010. Another conjecture to explain the spread is that quantify the directional spread of CM, we tested CM are an opportunistic, weedy species, competi- whether the pattern of CM invasion deviated signif- tively superior to other cnidarians such as corals, and icantly from a uniform circle surrounding the ship by 123 Managing an invasive corallimorph plotting and testing for circular uniformity and reflec- recorded every 10 m moving along each transect line, tive symmetry (Rao Jammalamadaka and Sen Gupta and the average depth recorded per transect line. This 2001). Areal coverage of CM was calculated by strategy gave four depth categories from deep to plotting the infestation and measuring the area of shallow (e.g. tape 1 deployed at 27.4 m = deep reef splined polygons surrounding the limits of mild (Score and tape 4 deployed onto the reef flat = shallow). The 1), moderate (Score 2), or severe (Score 3) infestation. presence/absence of CM was recorded every m (point The original spread of the CM around the wreck intercept) along each 30 transect giving a total of 124 was on the western reef shelf of Palmyra Atoll. points at each site. CM cover for each depth category However, evidence of CM on reef slopes to the north (n = 4) was pooled for the 5 sites. Average percent and south of the atoll prompted us to record depth CM cover for the four depth categories was compared distribution of CM on the forereef at five sites using Kruskall–Wallis analysis of variance, because surrounding the atoll (Fig. 1a). These sites were data did not fit assumptions of normality by the chosen because they are historically established ben- Shapiro–Wilks test (W = 0.68365, p = 2.48e-05). thic survey stations for researchers at Palmyra Atoll. At each site, transect lines were initiated at a depth of Oceanography 27.4 m and deployed perpendicular to shore up the reef slope and onto the reef flat to a depth of ca. 0.1 m. Once established, it was expected that CM spread from This configuration required four 30 m transects to be the shipwreck site would occur via the dispersal of laid end-to-end and separated by 1 m. Depth was larvae and clonal fragments directed by ocean Fig. 1 a Map of Palmyra showing the shipwreck site (circle) at 4 depth intervals: median percent corallimorph cover (line), and locations of depth transects (red triangles). Also shown are first and third quartile (boxes), whiskers (1.5 times the shallow reef flats (beige) and emergent land (green).