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Outbreak Densities of the Coral Predator Drupella in Relation to in Situ Acropora Growth Rates on Ningaloo Reef, Western Australia

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Outbreak Densities of the Coral Predator Drupella in Relation to in Situ Acropora Growth Rates on Ningaloo Reef, Western Australia Outbreak densities of the coral predator Drupella in relation to in situ Acropora growth rates on Ningaloo Reef, Western Australia C. Bessey, R. C. Babcock, D. P. Thomson & M. D. E. Haywood Coral Reefs Journal of the International Society for Reef Studies ISSN 0722-4028 Coral Reefs DOI 10.1007/s00338-018-01748-7 1 23 Your article is protected by copyright and all rights are held exclusively by Springer- Verlag GmbH Germany, part of Springer Nature. This e-offprint is for personal use only and shall not be self-archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Coral Reefs https://doi.org/10.1007/s00338-018-01748-7 REPORT Outbreak densities of the coral predator Drupella in relation to in situ Acropora growth rates on Ningaloo Reef, Western Australia 1,2 2,3 1 3 C. Bessey • R. C. Babcock • D. P. Thomson • M. D. E. Haywood Received: 25 September 2017 / Accepted: 26 October 2018 Ó Springer-Verlag GmbH Germany, part of Springer Nature 2018 Abstract Outbreaks of coral predators are defined as maximum number of Drupella that could be sustained increases (often rapid) in their abundance above threshold across a range of coral cover. Our data suggest that the densities that can be sustained by local coral assemblages, outbreak density of Drupella at the average level of coral which in turn depends on the abundance and turnover of cover for back reef sites at Mandu reef (17.6 ± 13.7%) is coral prey. To investigate the outbreak densities of the approximately 0.95 individuals m-2 reef area. At the corallivorous gastropod Drupella cornus, we conducted maximum coral cover observed at Mandu reef (60%), the both in situ feeding and coral growth experiments at outbreak density of Drupella is estimated to be approxi- Mandu reef within the Ningaloo Marine Park, Western mately 2.83 individuals m-2 reef area. Establishing Dru- Australia. Over two 10-day periods, we tagged and pho- pella outbreak densities assists managers in predicting tographed feeding scars on colonies of the tabulate coral possible outbreak abundances and in monitoring coral reef Acropora spicifera that harboured Drupella feeding health. aggregations. We calculated a mean in situ Drupella con- sumption rate of 1.16 ± 1.1 cm2 coral area individual-1 - Keywords Ningaloo Marine Park Á Western Australia Á d-1. We also determined coral growth rates by tagging and Management Á Coral growth Á Consumption rate photographing 24 colonies of Acropora spicifera at time zero and then again 1 year later. We calculated a mean linear extension rate of 7.9 ± 3.7 cm yr-1 for actively Introduction growing Acropora spicifera, which we then used to esti- mate A. spicifera growth rates over a range of coral cover Drupella is a genus of marine gastropods (Family Muri- values. This combination allowed us to determine the cidae) that feeds almost exclusively on the living tissue of corals (Robertson 1970; Claremont et al. 2011). These generalist corallivores occur throughout the shallow waters Topic Editor Morgan S. Pratchett of the Indo-Pacific (Claremont et al. 2011). However, excessive densities (outbreaks) of corallivores can result in & C. Bessey dramatic and widespread decline in coral cover (Pratchett [email protected] et al. 2014; Babcock et al. 2016). Outbreaks are defined as 1 Oceans and Atmosphere, Commonwealth Scientific and increases (often rapid) in the abundance of coral predators Industrial Research Organization, 64 Fairway, Crawley, above threshold densities that can be sustained by local WA 6009, Australia coral assemblages, which in turn depend on the abundance 2 School of Plant Biology and the Oceans Institute, University and turnover of coral prey. of Western Australia, 64 Fairway, Crawley, WA 6009, Outbreaks of the coral predator Drupella are associated Australia with high coral mortality (Moyer et al. 1982; Fujioka and 3 Oceans and Atmosphere, Queensland Biosciences Precinct, Yamazato 1983; Ayling and Ayling 1987; Turner 1994; Commonwealth Scientific and Industrial Research Organization, 306 Carmody Rd, St Lucia, QLD 4072, Antonius and Reigl Antonius and Riegl 1997; Shafir and Australia Gur 2008; Cumming 2009a). In Japan, coral degradation 123 Author's personal copy Coral Reefs was associated with large aggregations of Drupella, where relate to field monitoring data of coral cover. Furthermore, densities of Drupella spp. (mainly D. fragum) reached 5.12 little is known about in situ Drupella feeding rates. individuals m-2 (Moyer et al. 1982; Fujioka and Yamazato Growth of the preferred prey of Drupella is another 1983). In the Red Sea, densities of D. cornus were more essential component in calculating the outbreak densities of than 200 individuals per 30-cm-diameter coral colony on Drupella that can be sustained by the coral community. multiple genera, resulting in 100% coral mortality at the The growth rates of acroporids, the preferred prey of study site; the southern end of a 150-m artificial limestone Drupella, can vary depending on growth form, species, and quay (Shafir and Gur 2008). Similarly, correlations local environment, as well as the method used to quantify between declining coral cover and increasing frequency of their growth (Pratchett et al. 2015; Drury et al. 2017). D. cornus had also been observed in the Red Sea by Obtaining a measure of growth requires direct quantifica- Antonius and Riegl (1997), who reported average densities tion of colony linear dimensions, area, volume, or weight, of up to 12.24 Drupella m-2 (Al-Moghrabi 1997). On taken at various time increments to calculate a time-aver- Ningaloo Reef, in Western Australia, D. cornus densities of aged rate of growth (Pratchett et al. 2015). Linear exten- up to 19.4 m-2 have been associated with a 75% reduction sion, measured as a change in colony radius over a year, is in coral cover (Ayling and Ayling 1987; Turner 1994). a common metric of coral growth. There are relatively few Corals form the basic reef structure that provides habitat studies of coral growth rates in Western Australia. Annual for a myriad of organisms, including economically linear extension rates for Acropora spicifera are reported as important, iconic, and endangered species. Understanding 12.4 ± 1.4 cm at Ningaloo Reef (western side of the North the densities of Drupella that can be sustained based on West Cape) and 10.5 ± 1.2 cm at Bundegi Reef, which is coral cover and growth is essential information required to located within the Exmouth Gulf on the eastern side of the help managers monitor and predict possible outbreak North West Cape (Stimson 1996). Annual coral growth abundances. ranged from 7.3 to 14.6 cm for the closely related species Drupella feeding rates are a key component in calcu- Acropora hyacinthus in the Dampier Archipelago (Simp- lating the outbreak densities of Drupella that can be sus- son 1988). Ideally this growth should be expressed as tained by a coral community. Although Drupella can projected surface area, since photographic monitoring of forage on a range of coral morphologies (branching, tab- coral typically uses percent cover (e.g. Speed et al. 2013). ulate, massive and encrusting) and genera (Acropora, While linear extension rates may be a more accurate rep- Astreopora, Fungia, Millepora, Montipora, Pavona, resentation of colony growth, these are difficult to deter- Pocillopora, Porites, Seriatopora, Stylophora, and mine outside of a controlled laboratory environment and Turbinaria) using a specialised radula for scraping (Fujioka are not typically used in large-scale monitoring and Yamazato 1983; Ayling 2000; Shafir and Gur 2008; programmes. Hoeksema et al. 2013), they display a strong preference for The goal of our study was to determine the outbreak the acroporids (mainly Acropora and Montipora; Boucher densities of Drupella that can be sustained by a coral 1986; Turner 1994; Cumming 1999; Schoepf et al. 2010; community, based on coral cover and growth, in order to Moerland et al. 2016). Drupella tend to create discrete help managers predict possible outbreak abundances. We feeding scars and feed at the interface between live and conducted our study in the Ningaloo Marine Park, Western dead corallites during both day and night (although pre- Australia, where previously reported Drupella outbreaks dominantly at night) and show an attraction to stressed or have been associated with substantial declines in coral damaged coral (Forde 1992; Turner 1994; Morton et al. cover (Turner 1994). The purpose of our study was to (1) 2002; Bruckner et al. 2017). Field experiments report D. determine the percent cover of Drupella prey (hard coral cornus moving more than 2 m overnight to aggregate on and preferred prey species Acropora spicifera), (2) deter- damaged coral, potentially detecting prey species through mine Drupella density, (3) determine in situ Drupella species-specific chemical substances (Kohn 1961; Turner feeding rates on A. spicifera, (4) measure in situ growth 1994; Kita et al. 2005). Aquarium experiments estimate D. rates of A. spicifera, and (5) develop estimates of the cornus feeding rates at 2.3 cm2 of coral individual-1 d-1, maximum sustainable density of Drupella for maintenance where the average-sized Drupella was 28–35 mm (Cum- of net coral growth over a range of coral cover, thereby ming 2009b) and coral tissue was measured as the total quantifying a Drupella outbreak density for Ningaloo Reef.
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