Coral-associated invertebrates: diversity, ecological importance and vulnerability to disturbance

Pat Hutchings; Jessica Stella

The biodiversity of coral reefs is dominated by invertebrates. Many of these invertebrates live in close association with scleractinian corals, relying on corals for food, habitat or settlement cues. Given their strong dependence on corals, it is of great concern that our knowledge of coral-associated invertebrates is so limited, especially in light of severe and ongoing degradation of coral reef habitats and the potential for species extinctions. This review examines the taxonomic extent of coral-associated invertebrates, the levels of dependence on coral hosts, the nature of associations between invertebrates and corals, and the factors that threaten coral-associated invertebrates now and in the future. There are at least 860 invertebrate species that have been described as coral associated, of which 310 are decapod crustaceans. Over half of coral-associated invertebrates appear to have an obligate dependence on live corals. Many exhibit a high degree of preference for one or two coral species, with species in the genera Pocillopora, Acropora and Stylophora commonly preferred. This level of habitat specialization may place coral-associated invertebrates at a great risk of extinction, particularly because preferred coral genera are those most susceptible to coral bleaching and mortality. In turn, many corals are also reliant on the services of particular invertebrates, leading to strong feedbacks between abundance of corals and their associated invertebrates. The loss of even a few preferred coral taxa could lead to a substantial decline in invertebrate biodiversity and have far-reaching effects on coral reef ecosystem function. A full appreciation of the consequences of further coral reef degradation for invertebrate biodiversity awaits a more complete description of the diversity of coral-associated invertebrates, the roles they play in coral reef ecosystems, their contribution to reef resilience and their conservation needs.

Oceanography and Marine Biology: An Annual Review, 2011, 49, 43–104 © R. N. Gibson, R. J. A. Atkinson, J. D. M. Gordon, I. P. Smith and D. J. Hughes, Editors Taylor & Francis CoRAl-ASSoCIATED INvERTEbRATES: DIvERSITy, EColoGICAl IMPoRTANCE AND vulNERAbIlITy To DISTuRbANCE JESSICA S. STEllA1,2,3, MoRGAN S. PRATCHETT2, PAT A. HuTCHINGS4 & GEoFFREy P. JoNES1,2 1School of Marine and Tropical Biology, James Cook University, Townsville, Queensland 4811, Australia E-mail: Jessica.Stella@my.jcu.edu.au (corresponding author) 2ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, Queensland 4811, Australia 3 Climate Adaptation Flagship, CSIRO, Hobart, Tasmania 7001, Australia 4 The Australian Museum, Sydney, New South Wales 2010, Australia Abstract The biodiversity of coral reefs is dominated by invertebrates. Many of these invertebrates live in close association with scleractinian corals, relying on corals for food, habitat or settlement cues. Given their strong dependence on corals, it is of great concern that our knowledge of coralassociated invertebrates is so limited, especially in light of severe and ongoing degradation of coral reef habitats and the potential for species extinctions. This review examines the taxonomic extent of coral-associated invertebrates, the levels of dependence on coral hosts, the nature of associations between invertebrates and corals, and the factors that threaten coral-associated invertebrates now and in the future. There are at least 860 invertebrate species that have been described as coral associated, of which 310 are decapod crustaceans. over half of coral-associated invertebrates appear to have an obligate dependence on live corals. Many exhibit a high degree of preference for one or two coral species, with species in the genera Pocillopora, Acropora and Stylophora commonly preferred. This level of habitat specialization may place coral-associated invertebrates at a great risk of extinction, particularly because preferred coral genera are those most susceptible to coral bleaching and mortality. In turn, many corals are also reliant on the services of particular invertebrates, leading to strong feedbacks between abundance of corals and their associated invertebrates. The loss of even a few preferred coral taxa could lead to a substantial decline in invertebrate biodiversity and have far-reaching effects on coral reef ecosystem function. A full appreciation of the consequences of further coral reef degradation for invertebrate biodiversity awaits a more complete description of the diversity of coral-associated invertebrates, the roles they play in coral reef ecosystems, their contribution to reef resilience and their conservation needs. Introduction Coral reefs are complex and productive ecosystems that encompass the highest biodiversity of any marine ecosystem (Sebens 1994, Gray 1997, Hoegh-Guldberg 1999, veron 2000). Estimates of the number of species found on coral reefs range from 172,000 to over 9 million (Reaka-Kudla 1997, 43 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Ruppert et al. 2004). The uncertainty in species estimates is largely because like most marine ecosystems, coral reef biodiversity is dominated by highly diverse invertebrate taxa that are understudied and incompletely described (Reaka-Kudla 1997). The literature on coral reef organisms and their taxonomy has a strong bias towards the most conspicuous reef organisms, such as corals and ishes, whereas smaller, cryptic organisms remain largely ignored (Gaston 1991, bouchet et al. 2002). The estimates of reef ish and coral diversity stand at about 4000 (Choat & bellwood 1991, lieske & Myers 1994, bellwood et al. 2003) and about 800 species, respectively (Paulay 1997, veron 2000, Hughes et al. 2002). However, invertebrates other than corals account for the vast majority of animal species on coral reefs. It is an unfortunate circumstance that, given the documented global threats to coral reef ecosystems (Sebens 1994, Hoegh-Guldberg 1999, Gardner et al. 2003, Hughes et al. 2003, bellwood et al. 2006), the taxa that account for the greatest biodiversity are those that have received the least attention. The extraordinary biodiversity of coral reefs is evident at all taxonomic levels (Table 1). Members of all the 34 extant animal phyla, except the onycophora and Cycliophora, occur on coral reefs (Castro 1988, Gray 1997, Williamson 1997, Ponder et al. 2002, Ruppert et al. 2004). The most recent estimates indicate that there are at least 165,000 described invertebrate species associated with coral reefs (Table 1). In terms of described species, the fauna is dominated by molluscs, arthropods, nematodes, platyhelminthes and cnidarians. The true diversity of coral reef invertebrate species will not be known until there are many more systematic studies of most groups (but see Abele 1974, Austin et al. 1980, Edwards & Emberton 1980, black & Prince 1983). It is estimated that 1635 new marine species are currently described every year, mostly crustaceans and molluscs (bouchet 2006), but sampling effort remains insuficient for most invertebrate groups (bouchet et al. 2002). Even without precise species counts, it is clear that invertebrates account for the greatest biodiversity on coral reefs, regardless of the taxonomic level by which it is measured (Ray 1985, Earle 1991, Ray & Grassle 1991, Williamson 1997, Ruppert et al. 2004). Despite their ubiquity, the factors affecting biodiversity and abundance of invertebrates on coral reefs are poorly understood. Consequently, understanding of coral reef processes is drawn from knowledge of a relatively small proportion and taxonomically biased selection of coral reef species. The high biodiversity of coral reef organisms is partly attributed to the extraordinary diversity of habitats and topographic complexity provided by scleractinian corals (luckhurst & luckhurst 1978, Sale 1991, McClanahan 1994, Jennings et al. 1996, Öhman & Rajasuriya 1998, lawson et al. 1999, lindahl et al. 2001, Gratwicke & Speight 2005, Garpe et al. 2006, Wilson et al. 2007). In addition, coral reefs have been associated with the evolution of the largest diversity of symbiotic associations in the marine environment (Castro 1988). Many of these symbioses involve branching corals and other reef invertebrates (Patton 1966, 1974, Tyler 1971, bruce 1972b, 1977, Abele & Patton 1976, Austin et al. 1980, Coles 1980, Chang et al. 1987, Castro 1988, Tsuchiya et al. 1992, Sin 1999, Stewart et al. 2006). Symbiotic associations with branching corals have many beneits for reef invertebrates; corals provide them with a large surface area on and in which to live, refuges from predation, food in the form of coral tissue, mucus and its associated detritus, and a hard skeleton used as a substratum by specialized burrowers and gall-forming animals (Castro 1988). In utilizing coral in such ways, many reef invertebrates have become reliant on the coral substratum. Whereas some utilize coral hosts indiscriminately, a high proportion of them may exhibit a degree of specialization to a living coral host, and this mode of life can often involve lifelong associations between individuals. The relationship coral-associated invertebrates have to corals can be either obligate (must live on their coral host to survive) or facultative (may live on a coral host but does not have to for survival) (Castro 1976). The difference between obligate and facultative symbiosis may have fundamentally different consequences for the structure and dynamics of these invertebrate assemblages. An understanding of the degree of dependence of invertebrates on corals has become a particularly important issue in light of the increasing variety of factors contributing to a global decline in coral cover (Sebens 1994, Hoegh-Guldberg 1999, Jones et al. 2004, bellwood et al. 2006). A 44 CoRAl-ASSoCIATED INvERTEbRATES Table 1 Distribution and species count estimates of invertebrate phyla by habitat Phylum onycophora Placozoa orthonectida Porifera Cnidaria Ctenophora Platyhelminthes Nemertea Gnathostomulida Rotifera Gastrotricha Kinorhyncha Nematoda Nematophora Priapulida Acanthocephala loricifera Annelida Echiura Sipuncula Mollusca Arthropoda Tardigrada brachiopoda bryozoa Phoronida Echinodermata Chaetognatha Hemichordata Dicyemia Kamptozoa Cylciophora Seisonida Chordata Total Marine Coral reefs Freshwater X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X Terrestrial Coral reef species X 0 1 20 7,800 9,980 80 20,000 1,120 80 X X X X X X X X X X X X X X X X X X X X X X X X 50 500 150 20,000 4 2 1,150 100 8,000 150 150 50,000 35,000 800 350 4,000 14 6,000 150 70 75 150 0 3 2,300 168,249 Source: updated, based on Ray & Grassle 1991 and Ruppert et al. 2004. decline in coral cover will presumably affect the animals that interact with living coral, and the nature of these interactions may be used to predict the effects of declining coral cover on coral associates. If particular coral species disappear, facultative associates may persist by switching hosts, whereas the more specialized, obligate associates will be at greater risk of extinction (McKinney 1997). The co-dependence between obligate associates and corals appears to be so strong that some species of corals do not seem able to persist without their symbionts (Glynn 1983, Stewart et al. 2006), and obligate associates are only found in association with certain coral hosts (Glynn 1983). understanding how specialized species are to a certain habitat is important because these factors can greatly enhance the risk of extirpation or extinction following declines in the availability of potential habitats (lawton 1993, McKinney 1997, Pratchett et al. 2008). Habitat degradation is a 45 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Number of species vs. number of studies (log transformed) 6 Number of species Number of studies 5 4 3 2 1 0 Invertebrates Fish (includes sharks and rays) Corals Coral reef taxa Figure 1 Number of bleaching-related studies conducted compared with the number of species of coral reef taxa. Species numbers were log transformed due to the disproportionately large number of invertebrate species. values based on search in ISI Web of Science database in June 2010. major threat to coral reefs worldwide, so it is imperative that we understand the degree of specialization and codependence of these symbiotic relationships and learn more about how disturbance will affect their diversity. Current research on the potential impact of coral bleaching on the biodiversity of reef organisms highlights the disparity in our knowledge of different taxonomic groups. To date, much of the focus has been on the impact of bleaching on the corals themselves and on the consequences for reef ishes (Figure 1). Despite the disproportionate diversity represented by invertebrates, there are practically no data on the coral-invertebrate relationship and the potential effects of coral reef degradation via prolonged and frequent bleaching or coral death on their biodiversity (Figure 1). Studies of coral-reef ish associations have documented a high degree of dependence on the coral substratum and the consequent dramatic effects of habitat loss and degradation on reef ish assemblages (e.g., Reese 1977, Harmelin-vivien & bouchon-Navaro 1983, Sano 1989, Hughes 1994, Shears & babcock 2002, bellwood et al. 2003, Duffy 2003, Hughes et al. 2003, Jones et al. 2004, Munday 2004, Garpe et al. 2006, Graham et al. 2006, Pratchett et al. 2006, Wilson et al. 2006). by comparison, the nature of the association between corals and other invertebrates and their response to human impacts on coral reef habitats are not well known even though their association may be stronger and the threat to reef biodiversity may be greater than for ishes. To assess the likely effects on reef biodiversity due to coral reef degradation, it is irst necessary to focus clearly on the largest component of reef biodiversity: the invertebrates. To date, there has not been a systematic review of all known coral-associated invertebrates and the coral taxa with which they associate. The aim of this review is to assemble and synthesize the disparate literature on the diversity and taxonomic composition of coral-associated invertebrates and their hosts. First, an assessment is made of the roles coral-associated invertebrates play in reef processes and the nature of their association with scleractinian corals, examining both positive and negative feedbacks in invertebrate-coral host relationships. Second, the patterns of habitat use and specialization are examined, including the extent of their coral dependence across a range of invertebrate groups for which information is now available. Finally, both the anthropogenic and 46 CoRAl-ASSoCIATED INvERTEbRATES environmental threats they are currently facing and how habitat specialization may enhance their risk of extinction are assessed. Biodiversity of invertebrates that associate with scleractinian corals There have been numerous isolated studies and anecdotal reports since the 1930s of invertebrates that associate with coral. Most published studies have focused on single taxonomic groups such as polychaetes and decapods, and relatively few have quantiied the diversity of all invertebrates found within select coral hosts (Patton 1966, 1974, Abele & Patton 1976, Austin et al. 1980, Coles 1980, Edwards & Emberton 1980, black & Prince 1983, Stella et al. 2010). The Appendix (see p. 83) lists all invertebrates known to associate with scleractinian coral, the species of coral with which they have been known to associate, any documented explanations of the association, and the level of coral dependence exhibited (i.e., whether obligate or facultative). A total of 211 published studies were included in this analysis, documenting an immense diversity of invertebrate species that associate with and may rely on coral for food, for a habitat or as a substratum on which they graze or a combination of all three. All names were carefully cross-checked amongst the literature for synonyms or name changes to ensure the most current and accurate assessment. We have identiied 869 invertebrate species found to associate with scleractinian corals. These species are distributed among 108 different families and eight phyla: Arthropoda, Mollusca, Echinodermata, Annelida, Porifera, Platyhelminthes, Sipuncula and Hemichordata (Appendix). The representation of species varied greatly among taxa, with a disproportionate number of species belonging to Phylum Arthropoda (Appendix). The total of 869 species included 636 arthropods, 130 molluscs, 51 echinoderms, 29 annelids, 11 poriferans, 9 platyhelminthes, 2 sipunculans, and 1 hemichordate (Appendix). of the 636 arthropods, 310 were decapod crustaceans, constituting 35% of the total number of invertebrate species to utilize coral (Appendix). The overwhelming contribution of arthropods to the overall diversity, particularly by decapod crustaceans, is a common theme in the literature (Abele 1974, 1976, Patton 1974, Abele & Patton 1976, Austin et al. 1980, Stella et al. 2010). Decapods appear to have the most intimate association with a coral host, as indicated by their heavy reliance on a host for food, habitat and reproduction and their adverse reactions to a decline in coral host health (Glynn et al. 1985, Tsuchiya et al. 1992). The high diversity of coral-associated invertebrates known thus far is likely to inluence coral reef processes by the numerous potential species interactions, not only amongst each other but also with their coral host. Invertebrates associate with coral for different reasons, and their association may be a brief encounter or a lifelong partnership. Moreover, some associates are free living, whereas others actually live with their coral host in a co-dependent symbiotic relationship. Whether an association is obligate or facultative has different consequences for associates should their host corals decline in health or disappear entirely. Invertebrates that utilize a coral host as a necessity for survival, whether for food or habitat, are recognized as obligate coral associates. A facultative associate, on the other hand, is one that may beneit from utilizing coral but can survive without it. of the 869 invertebrates reviewed, 56% (487) were found to be obligate coral associates (Figure 2). This is an unexpected and alarmingly high proportion because it is extremely unlikely that these animals would be able to persist should their coral host decline in health or die. because obligate species are reliant on their coral host for survival, their population dynamics and distribution may be inextricably linked with that of their host. Almost all obligate species are either arthropods (410) or molluscs (61) (Appendix). Although never before considered, severe loss of coral habitat, due to climate-induced coral bleaching and other factors causing reef degradation, is likely to cause localized extinctions among these coral-dwelling invertebrates, potentially causing massive declines in biodiversity. Thus, coral reef 47 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Number of species reviewed 500 56% 400 33% 300 200 11% 100 0 Obligate Unknown Facultative Nature of relationship with coral host Figure 2 Coral-associated invertebrates categorized by their level of dependence on a coral host. ecosystems are likely to experience a large proportional change in biodiversity as coral reef habitats are continually degraded. Also of concern are the 283 species for which knowledge of coral dependence is unknown. Without further studies, it is impossible to speculate into which category these species should be classiied. The reason why coral-associated invertebrates use a coral host is an important factor in determining how reliant on a host they are. Coral dwellers are those that utilize coral as habitat, refuge from predation as well as mating sites within the protection of coral branches (Castro 1988, Munday et al. 1997). Corallivores consume live coral tissue, mucus or coral skeleton. Corallivorous invertebrates can inlict minor or lethal damage on their coral hosts, which subsequently can have deleterious effects on coral growth and itness (Rotjan & lewis 2008). Invertebrate corallivores employ a wide range of feeding strategies and as such can be obligate or facultative coral feeders (Robertson 1970). The largest proportion of coral-associated invertebrates was found to utilize a coral host for habitat or food (Appendix, Figure 3). Habitat was the primary reason for use for 332 species, and 314 species have been documented to consume coral, with only 26 species using coral as a substratum (such as for grazing) (Figure 3). For the remaining portion of coral-associated invertebrates (197 species), there is no information on the reason for coral use, only a record documenting that the species is found associated with a coral host (Figure 3). It is also unknown whether some of these species also occur on non-coral hosts. The majority of arthropods utilized coral as habitat (282 species) and food (239 species) (Figure 3). Molluscs were the second-largest component of coral-associated invertebrates, with 34 species that utilize coral as habitat and 53 species that consume coral tissue (Figure 3). Classifying coral-associated invertebrates by coral use is not always straightforward. For instance, obligate species, such as trapeziid and tetraliid crabs and alpheid shrimp, use branching coral for both food and habitat, yet it is unclear which takes precedence. Relationships between invertebrates and their coral hosts Scleractinian corals have three important roles in the ecological function of coral reef ecosystems: (1) foundation species, contributing signiicantly to basal carbon production for complex reef-based food webs (Hatcher 1988); (2) habitat-forming species, providing critical habitat for coral-dwelling animals (e.g., Munday et al. 2001); and (3) structural engineers (Jones et al. 1994), contributing to high habitat complexity and surface topography, which promotes biodiversity by mediating competition or predation (Menge 1976, Holt 1987, Hixon & Menge 1991, Coker et al. 2009). High cover and diversity (e.g., growth forms) of corals are therefore fundamental in sustaining high diversity 48 CoRAl-ASSoCIATED INvERTEbRATES 700 Habitat 600 Food Unknown Number of species 500 Substratum 400 300 200 100 0 Ar p ro th od a s lu ol M ca od in m er at a a lid e nn A es a er rif Po h Ec Pl th in lm he y at Si p c un ul a ta da r ho ic m e H Phyla Figure 3 Patterns of coral use among coral-associated invertebrates by phyla (based on documented evidence). of coral-associated, as well as reef-associated, organisms (Pratchett et al. 2009). It is apparent, however, that there are numerous feedbacks in which the coral-associated organisms that depend on scleractinian corals are, in themselves, contributing to temporal and spatial patterns of abundance for scleractinian corals. Feedback mechanisms that reinforce high cover and diversity of corals may include the local production of critical nutrients (such as nitrogen and phosphorus) by diverse suites of coral-associated organisms, which may enhance coral growth (sensu Meyer & Schultz 1985). Conversely, coral-feeding invertebrates, especially those species that occur at very high densities (e.g., outbreaks of Drupella snails, and Acanthaster starishes), can destabilize temporal dynamics of coral abundance, contributing to rapid declines in coral cover (e.g., Chesher 1969, Pearson & Endean 1969, yamaguchi 1986). Negative feedback The most apparent effects of coral-associated invertebrates on corals are direct negative effects associated with coral feeding. However, coral feeding (by both ishes and invertebrates) is generally regarded to have negligible effects on corals (reviewed by Cole et al. 2008, Rotjan & lewis 2008). The main exception is those species that exhibit population explosions (termed ‘outbreaks’) and thereby cause extensive and widespread coral mortality. Much of the research on invertebrate corallivores has been directed towards those species with the potential to cause rapid and extensive coral loss, such as the crown-of-thorns starishes Acanthaster spp., (Glynn 1974, Glynn & Krupp 1986, Williams 1986, Glynn 1994, Reyes-bonilla & Calderon-Aguilera 1999, Pratchett 2005) and the gastropod snails Drupella spp. (boucher 1986, Turner 1994, McClanahan 1997) and Coralliophila spp. (brawley & Adey 1982, baums et al. 2003a,b). There is, however, an exceptional, and ever-growing, list of invertebrate species known to feed on scleractinian corals. Extensive searches of published 49 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES sources revealed 314 invertebrate species, from 24 families and 5 phyla, that have been reported to feed on scleractinian corals (Figure 3). This greatly exceeds the number of vertebrate species (128 species) known to feed on scleractinian corals (Cole et al. 2008) and is much higher than previous estimates of the diversity (51 species) of invertebrate corallivores (Rotjan & lewis 2008). of the 314 corallivorous species revealed during this study, the vast majority (76%, 239 species) were arthropods (Appendix). The remaining species were molluscs (53 species), echinoderms (12 species), platyhelminthes (9 species) and polychaetes (1 species) (Appendix). The high diversity of arthropods that consume coral included many species of obligate coral-dwelling decapod crabs (e.g., Cymo, Trapezia and Tetralia), and endoparasitic copepods (Corallovexia, Xariia and Corallonoxia). obligate coral-dwelling crabs are often observed grazing on their host corals, feeding on either live tissue, mucus or fat bodies produced by Pocillopora corals (e.g., Knudsen 1967, Stimson 1990, Rinkevich et al. 1991, Castro et al. 2004). Given their small size and generally low densities on each coral colony (typically ≤ 2 individuals per colony (e.g., Stella et al. 2010) it is often assumed that these common and widespread coral crabs do not have any signiicant negative effects on their hosts (Rotjan & lewis 2008). Moreover, some species (e.g., Tetralia) feed predominantly on mucus rather than live tissue (Stimson 1990), which could have signiicant beneicial effects (Stewart et al. 2006), as discussed further in this chapter. Impacts of most corallivorous organisms (including many polyp-feeding and mucus-feeding ishes) are presumed to be negligible (e.g., Harmelin-vivien & bouchon-Navaro 1983, Hixon 1997) because they cause little apparent damage to prey corals (Rotjan & lewis 2008). Recent research on effects of corallivorous ishes on prey corals revealed that energetic costs of chronic predation could be signiicant (e.g., Cole et al. 2009), and that differential feeding among coral species could inluence the relative abundance of preferred versus non-preferred prey (bellwood et al. 2006, Cole et al. 2010). Effects of coral-feeding ishes may be particularly pronounced following extensive depletion of prey (e.g., after mass bleaching) whereby feeding is concentrated on the few remaining corals and may represent the ultimate cause of coral mortality (bellwood et al. 2006). Similarly, obligate coral-dwelling crabs have speciic associations with a select number of different corals and, together with diverse assemblages of other coral-feeding organisms or in conjunction with other coral disturbances, may contribute to selective coral mortality. In the Chagos Archipelago, central Indian ocean, high densities of the decapod Cymo melanodactylus (up to 47 crabs per colony) were recorded on colonies of Acropora cytherea that exhibited extensive areas of recent mortality (Pratchett et al. 2010). It is unknown whether these coral-feeding crabs caused, or contributed to, observed coral mortality, but more research is needed to assess the effects of chronic predation by small, persistent, coral-associated invertebrates. The greatest diversity of coral-feeding invertebrates is represented by parasitic copepods. At least 243 species of copepods are known to associate with live coral, of which 199 species parasitize their coral host (Appendix). Although their typical body size is about 1 mm, copepods may occur in high number; up to 668 individual copepods have been recorded within a single (ca. 16-cm diameter) colony of Pocillopora damicornis (Humes 1994), although it is unknown what effect, if any, they have on the growth, reproduction or survivorship of heavily infested corals (Humes 1985a). It is assumed that most copepods feed on coral mucus (especially endoparasitic species) rather than live tissue (Humes 1985a). However, recently Cheng & Dai (2010) analysed the gut contents of a copepod, Xariia issilis, and found an abundance of unicellular algae with characteristics of Symbiodinium, which they obtained from their coral host, Pocillopora damicornis. Interestingly, the algal cells remained viable in the gut after 2 weeks of starvation in the laboratory. Some copepods were also retained in either an all-light or an all-dark environment. After 5 days, all copepods in the dark treatment died, possibly due to starvation because photosynthesis would have stopped. because the algal cells remained photosynthetically active for a certain period of time, it was suggested that Xariia issilis may temporarily retain the ingested Symbiodinium, possibly for the beneits obtained through the release of photosynthetic products to the hosts (Cheng & Dai 2010). The 50 CoRAl-ASSoCIATED INvERTEbRATES use of obtained Symbiodinium by marine invertebrates has been widely documented (Trench & Winsor 1987, Trench 1993, lobban 2002, barneah et al. 2007), but this situation has yet to be tested for a comprehensive range of coral-associated species. Crown-of-thorns starish (nominally Acanthaster planci, but see vogler et al. 2008) are among the best-studied invertebrate corallivores and are renowned for their capacity to devastate local coral assemblages (see reviews by Potts 1981, Moran 1986, birkeland & lucus 1990). ordinarily, crown-of-thorns starish occur at very low densities (typically < 1 starish ha−1) and have little effect on the abundance of reef corals (e.g., Glynn 1973, Zann et al. 1990). However, each starish consumes up to 6 m2 of live coral per year (birkeland 1989), and the cumulative impacts of highdensity populations (up to 20,000 ha−1) can result in extensive and widespread coral loss (Carpenter 1997). Consequently, there has been considerable research focused on understanding when and why outbreaks occur. The explanation that has the greatest traction at present relates to increased nutrient loads in tropical waters or enhanced phytoplankton levels (brodie et al. 2005, Houk et al. 2007, Fabricius et al. 2010) because increased abundance of planktonic prey increases survival rates of larval A. planci (Fabricius et al. 2010), leading to increases in local adult abundance 3 years thereafter (birkeland 1982). Conversely, the removal of predatory ishes may increase survival and abundance of postsettlement starishes, resulting in increased incidence of outbreaks in areas with higher ishing pressure (Dulvy et al. 2004, Sweatman 2008). There is, however, little evidence that postsettlement stages of A. planci are actually prone to predation (Sweatman 1995). ultimately, crown-of-thorns starish are predisposed to major population luctuations due to inherent properties of their life history, such as immense fecundity, density-dependent fertilization successes, and short generation times (Moore 1978, Stump 1992). Furthermore, it is likely that there are multiple causes for outbreaks (Pratchett 2005), which may also vary regionally. outbreak densities of A. planci have been reported throughout the Indo-Paciic (Moran 1986). However, destructive effects of A. planci have been mostly restricted to the western Paciic, especially southern Japan, Australia’s Great barrier Reef, Guam and Fiji (Moran 1986, birkeland & lucas 1990, bruno &Selig 2007). Differential impacts of A. planci outbreaks in different geographic areas may relate to regional variation in the structure of coral communities (Pratchett 2010), whereby the greatest coral loss tends occur in areas dominated by Acropora, which is the preferred coral prey for Acanthaster planci (De’ath & Moran 1998, Pratchett 2001, 2007). Crown-of-thorns starish are well adapted to feeding on a wide range of different corals but often exhibit a striking preference for a small suite of the available prey species, which causes differential mortality among coral species, and can exert a major inluence on coral community structure (Pratchett 2001). In the eastern Paciic, Glynn (1974, 1976) found crown-of-thorns starish fed mostly on rarer coral species, increasing the dominance of the abundant coral species, Pocillopora damicornis (see also branham et al. 1971). Elsewhere, Acanthaster planci tend to feed mostly on relatively abundant coral species (e.g., Acropora and Montipora) and thereby increase the prevalence of non-preferred corals (e.g., ormond et al. 1976, Colgan 1987, Keesing 1992, De’ath & Moran 1998). Aside from feeding on prey corals, invertebrate corallivores have been implicated in transmitting or increasing vulnerability to coral disease (e.g., Sussman et al. 2003, Nugues & bak 2009), which indirectly contributes to coral loss or shifts in community composition. Controlled experimental studies have been done that suggest that the corallivorous ireworm Hermodice carunculata (Sussman et al. 2003), the gastropod Coralliophila abbreviata (Williams & Miller 2005) and a corallivorous nudibranch, Phestilla sp. (Dalton & Godwin 2006), are effective vectors for coral disease. There are further anecdotal observations suggesting that Drupella cornus (Antonius & Riegl 1997a,b) and Acanthaster planci (Nugues & bak 2009) may also contribute to spread of coral diseases. The ireworm Hermodice carunculata plays an important role as a winter reservoir and subsequent vector for the bacterium Vibrio shiloi, which causes bleaching of Oculina patagonica in the Mediterranean (Sussman et al. 2003). Initial instances of Vibrio shiloi infections at the start of summer correspond with the distribution of Hermodice carunculata, although there is also potential 51 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES for subsequent localized transmission independent of the ireworm (Sussman et al. 2003). In the Caribbean, Williams & Miller (2005) showed that the gastropod Coralliophila abbreviata effectively transmits disease (potentially white pox: Rodríguez-Martínez et al. 2001) among colonies of Acropora cervicornis when sequentially feeding on infected and uninfected colonies. In contrast, Coralliophila abbreviata snails taken only from uninfected colonies and allowed to feed on control colonies did not cause any disease infections (Williams & Miller 2005), suggesting that transmission potential may be short lived. In contrast, experimental studies using Phestilla sp. revealed that that after feeding on infected coral tissue the same individual nudibranch could sequentially infect ive different fragments of Turbinaria mesenterina over 12 days (Dalton & Godwin 2006). Moreover, corals infected with a tissue-sloughing disease after being consumed by Phestilla sp. were contagious and could be used to reinfect further coral fragments through direct contact. The agent of this disease was not conirmed but is suspected to be a Beggiatoa bacterium (Dalton & Godwin 2006). In Indonesia, Nugues & bak (2009) found that several (ive of eight) colonies of Acropora cytherea that had been partially consumed by Acanthaster planci were secondarily affected by brown-band disease. These results suggest that A. planci may be a vector of coral disease that transmits primary pathogens during feeding (Nugues & bak 2009). Alternatively, the starish may simply facilitate the spread of the disease, whereby areas of tissue loss provide an entry point for disease pathogens to infect additional coral colonies (e.g., Page & Willis 2007). Similarly, an outbreak of ‘white syndrome’ among Acropora corals in the Red Sea was correlated with an outbreak of Drupella cornus (Antonius & Riegl 1997a). It was unclear, however, whether coral mortality caused by white syndrome attracted or beneitted D. cornus or whether feeding activities of highdensity populations of the corallivorous snails contributed to the disease epidemic (Antonius & Riegl 1997a). At least two different coral diseases (brown-band disease, Aeby & Santavy 2006; skeletal-eroding band, Page & Willis 2007) have been shown to readily and rapidly infect coral fragments that were subject to experimentally induced injuries, whereas comparable coral fragments with intact coral tissue were almost never infected. In each of these experiments, injuries were imposed by air blasting or physically scraping the live coral to expose small areas (1–4 cm2) of underlying skeleton (Aeby & Santavy 2006, Page & Willis 2007). As such, any corallivores that remove coral tissue and expose the underlying skeleton, including Acanthaster planci and Drupella spp., may predispose corals to disease. Injuries to live corals inlicted by coral-feeding organisms may also facilitate the establishment of epibionts (e.g., Spirobranchus giganteus) and allow boring organisms to access the skeleton of live corals (Hutchings 1986). Many different sessile invertebrates, such as polychaetes and molluscs, live on or within live corals (Hutchings 1986), although their effects on host corals are generally unknown. only a few speciic coral-epibiont associations have ever been studied, such as interactions involving Spirobranchus polychaetes and vermetid gastropods. Although epibionts clearly occupy space that might otherwise be illed with coral polyps, Strathmann et al. (1984) and Devantier et al. (1986) suggested that Spirobranchus spp. generally have beneicial effects for host corals (as discussed in the next section). vermetid gastropods, however, feed with extruded mucus nets that can smother surrounding substrata, including corals (Colgan 1985). In French Polynesia, Shima et al. (2010) showed that colonization by the vermetid gastropods Dendropoma maximum, reduced skeletal growth of host corals by up to 81% and reduced survival by up to 52%. vermetid gastropods also had a disproportionate effect on Pocillopora corals and may therefore contribute to increased abundance of other corals (mostly Porites) within local coral assemblages (Shima et al. 2010). Similarly, some other sessile coral-associated invertebrates, such as sponges, may directly compete with host corals for common resources (Suchanek et al. 1983, Aerts & van Soest 1997, Rützler 2002). Sponges are able to damage corals by producing active substances even without direct contact (Suchanek et al. 1983, Sullivan et al. 1983, Porter & Targett 1988). Thus, competitive interactions between sponges and corals can often result in the overgrowth or death of the coral 52 CoRAl-ASSoCIATED INvERTEbRATES (Macintyre et al. 2000, Rützler 2002), although, as discussed in the next section, some sponges are mutualistic (Goreau & Hartman 1963). bioeroding activities of many boring organisms can be highly detrimental for host corals, undermining their structural integrity, making them more susceptible to being dislodged during storm events (Hutchings 2011) or potentially leading to structural collapse (Goreau & Hartman 1963). Most commonly, macroboring coral-associated animals are polychaete worms, sipunculans, bivalve molluscs and sponges (Hutchings 2011). Although a few boring species, such as barnacles, occur in living coral the majority avoid live coral due to the high predation of settling larvae by the corals (Hutchings 2011). The majority of larvae of boring organisms, therefore, settle on dead or damaged parts of the coral colony, typically at the base of the coral head. boring sponges are the best studied, and most signiicant of all boring animals (Hutchings 1986), accounting for up to 94% of skeletal excavation. However, boring sponges are relatively slow colonizers. Rather, it is the polychaetes that are the initial colonizers of newly available coral substrata, and they may actually facilitate subsequent colonization of boring sponges and sipunculans (Hutchings & bamber 1985, Pari et al. 2002). once these borers have settled and bored into the substratum they are effectively entombed for the rest of their life. Each borer produces characteristic burrows that can be identiied (Hutchings 1986, 2008). These burrows, when vacated by the death of the borer, provide favourable habitat for a wide range of other non-boring species, commonly referred to as nestlers or cryptofauna (Hutchings 1986), including many species that only occur on coral reefs. bioerosion may create an extensive three-dimensional structure, which may then be inilled by sediment and lithiication, thus strengthening the habitat (Wilkinson 1983). The process of bioerosion is generally most pronounced in the aftermath of coral death, whereby erect coral colonies may be reduced to rubble within 4–6 years (Sheppard et al. 2002, Graham et al. 2006). Internal bioeroders may be abundant, especially in areas enriched with nutrients (e.g., Fabricius 2005), and thus can greatly compromise coral growth, if not survival. Positive feedback Despite the large number of coral-associated invertebrates that directly feed on live corals, or otherwise contribute to reduced health and mortality of corals, there are certain species (e.g., obligate coral-dwelling crabs) considered to be fundamental to the persistence and resilience of their host corals (Glynn 1983). obligate coral-dwelling crabs from the family Trapeziidae, for example, are considered highly beneicial for host corals for two reasons: (1) they actively defend their host corals from larger and potentially devastating coral-feeding organisms, such as Acanthaster planci and Drupella cornus (Glynn 1982, 1987, vannini 1985, Pratchett et al. 2000, Pratchett 2001); and (2) in turbid conditions, they contribute to the removal of excess sediment, which can otherwise smother corals (Stewart et al. 2006). obligate coral crabs occupy virtually all branching species of both Acropora and pocilloporids, including Pocillopora, Stylophora and Seriatopora (Abele & Patton 1976, Stella et al. 2010), but the associates of pocilloporid corals are the most effective in repelling Acanthaster planci from feeding on host corals (Pratchett 2001). Differences in the symbiont assemblages of Acropora and pocilloporids are consistent across a wide range of coral species (Knudsen 1967, Tsuchiya & yonaha 1992). Most notably, Acropora species always contain Tetralia crabs, whereas pocilloporids always contain Trapezia species (Abele &Patton 1976, Patton 1994). The larger size (of both the carapace and chelipeds) of Trapezia species, compared with Tetralia crabs, may account for their increased eficacy in repelling Acanthaster planci (Glynn 1987). Moreover, behavioural observations have revealed that Trapezia crabs often attack the thorns of the starish, breaking them off at the pedicel, whereas Tetralia pinch mainly at the tube feet and, unlike Trapezia, do not cause any lasting damage to the starish (Glynn 1982, Pratchett et al. 2000). Glynn (1983) observed that these coral associates could detect approaching Acanthaster planci from a distance and began to exhibit a variety of 53 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES aggressive responses. Defence of host corals from A. planci may also be aided by shrimps (Alpheus and Coralliocaris) or coral gobies (Gobiodon and Paragobiodon), which alert crabs to any potential intruders or corallivores (see also vannini 1985). Pratchett (2001) demonstrated that the occurrence of coral-associated crustaceans (especially Trapezia and Alpheus) signiicantly inluenced the feeding preferences of Acanthaster planci. When given a choice of acroporid and pocilloporid corals with associates present, A. planci consistently selected acroporid corals. However, when all associates were removed the corals were consumed without selectivity. As such, widely reported feeding preferences of A. planci for Acropora over pocilloporid corals (brauer et al. 1970, Collins 1975, ormond et al. 1976, De’ath & Moran 1998) may relate to effectiveness of respective coral associates in defending their coral hosts. The scallop Pedum spondyloideum also repels Acanthaster planci from its coral host, massive Porites, using expellent water jets (Devantier & Endean 1988). Similarly, the rapid retraction of tube-forming polychaetes (e.g., Spirobranchus giganteus) tends to deter Acanthaster planci, but rather than preventing A. planci from eating their host colony, these organisms enhance the survival of only a few adjacent coral polyps, which may enable subsequent regeneration of the colony (Devantier et al. 1986, Devantier & Endean 1988). obligate coral-dwelling crabs further contribute to the health of host corals by actively removing sediment deposits from coral surfaces that would otherwise be detrimental to the health of the coral (Stewart et al. 2006). When sediments were experimentally added to host corals, crabs contained within colonies of both Acropora and Pocillopora “became highly active and began removing particles by ‘kicking’ with back appendages and ‘shoveling’ and ‘throwing’ them off with their chelae” (Stewart et al. 2006), which may contribute to increased survival of these corals in habitats with high-sediment regimes. To test this suggestion, Stewart et al. (2006) removed the obligate coral associates from healthy Acropora and Pocillopora corals located inside the lagoon in Moorea, French Polynesia. After 24 days, mortality rates for acroporid and pocilloporid colonies divested of associates were 45% and 80%, respectively. Whole-colony mortality resulted from excessive sedimentation and subsequent bleaching (Stewart et al. 2006). In contrast, nearby corals that retained intact assemblages of associates were healthy throughout. In the eastern Paciic, Glynn (1983) also showed that pocilloporid corals occupied by Trapezia crabs, as well as Alpheus shrimps, had a higher survival compared with corals divested of their crustacean associates. After 3 months, 31% of colonies without crustaceans experienced massive tissue loss. In contrast, corals occupied by crustaceans experienced no mortality and produced 19% more mucus than those without crustacean associates. Glynn (1983) argued that additional mucus production, stimulated by the presence of crabs, may protect the coral tissue from settling microorganisms, bacterial infections, sediment and invading larvae. The Pocillopora colonies harbouring crabs and shrimp demonstrated higher survival and growth rates than those deprived of crustaceans. on temperate reefs in North Carolina, Stachowicz & Hay (1999) observed a unique mutualistic association between an Oculina coral and the decapod crustacean, Mithraculus forceps. The crab relies on the coral for habitat and food in the form of lipid-rich mucus. In return, the crab defends its host from being overgrown by chemically noxious seaweeds like Dictyota and Sargassum that are avoided by most local herbivores. Corals from which crabs were experimentally removed developed a dense cover of epibionts and exhibited reduced growth and increased mortality relative to corals with crabs, which remained free of all epibionts. This association acts to promote the persistence of both species in habitats from which they might otherwise be excluded by competition and predation (Stachowicz & Hay 1999). Many obligate associates, such as trapeziid crabs, are trophically reliant on their coral host, using the coral tissue, mucus and associated detritus as their primary food source (Knudsen 1967, Glynn 1983). Trapezia live in symbiosis with their coral host, using it as both a food source and a refuge from predation. Although living within the branches of a coral host offers protection, these crabs are potential prey for other reef organisms. Trapeziid crabs are consumed by a number of reef 54 CoRAl-ASSoCIATED INvERTEbRATES ishes, including squirrel ishes, lounders, wrasse, moray eels, hawkishes and sweepers (Hiatt & Strasburg 1960). Trapeziid crabs move amongst coral colonies at night (Castro 1978), and there is some opportunistic generalized predation on these crabs, but most ishes are not able to prey on the crabs while the crabs are within the protection of their coral host. However, the wrasse Gomphosus varius specializes in foraging between coral branches, using a slender, protruding snout as forceps to extract crustaceans from between coral branches (Hiatt & Strasburg 1960, Hobson 1974, Sano et al. 1984). Hobson (1974) found that coral-associated crustaceans formed up to 70% of the wrasse’s diet in Hawaii. Coral associates, such as Trapezia, are therefore part of a multilevel trophic system-energy low through both horizontal and vertical pathways stemming from their coral host and extending to the wider-ranging food web (Rinkevich et al. 1991). Importantly, obligate coral associates may actively participate in nutrient recycling, whereby their excretion and excrement may enrich nutrient availability within the local vicinity of coral polyps (Patton 1976). Conirmation of nutrient enrichment was shown in the interaction between the shrimp Periclimenes yucatanicus and its anemone host Condylactis gigantea (Spotte 1996). The shrimp excretes ammonia, which consequently enriches the nitrogen concentration of water surrounding the tentacles of the anemone. Spotte (1996) found that anemones associated with the shrimp had a greater ability to take up external ammonia and contained a greater concentration of zooxanthellae within their tissue than those without the symbiont. Similarly, the mytilid bivalve Lithophaga simplex, which commonly inhabits Astreopora myriophthalma in the Red Sea, has been shown to enhance ammonium contributions, which beneit the host coral (Mokady et al. 1998). Lithophaga simplex, which bores into the skeletons of living coral colonies, is usually considered to be parasitic in association with corals. This view of the bivalves is based on the ‘obvious’ damage associated with bioerosion of the coral skeleton. However, the beneit provided through nutrient enrichment may signiicantly outweigh the cost of localized structural damage (Mokady et al. 1998), so caution must be exercised in ascribing the nature of interactions (parasitic or mutualistic) between corals and their resident associates. In contrast to the damage boring sponges can cause to corals, some sponges can actually prevent coral heads from being dislodged, due to the added lexibility of the basal part of the coral colony, which allows the colony to bend with the water currents (Goreau & Hartman 1963). Filter-feeders, such as sponges, have been found to play a crucial role in nutrient and carbon cycling on the reef, and those that live in association with coral may be particularly important in improving water quality and transporting nutrients into the vicinity of the coral colony (Richter et al. 2001, Ribes et al. 2005). The ability of some associates to protect corals from predators, alleviate detrimental effects of sedimentation, inhibit algal overgrowth and enrich nutrient concentration has important implications for the persistence of coral reef ecosystems. These abilities suggest that some associates increase coral vitality and may play a key role in coral resiliency by providing services beneicial to coral host health. For sessile corals, with limited innate defences, their symbiotic relationships with other invertebrates may be vital to the persistence and population dynamics of the coral. Patterns of coral use and specialization Habitat availability has been shown to inluence the abundance and diversity of some coral reef invertebrates (Kohn & leviten 1976). Therefore, the local distribution and abundance of reef organisms will be positively correlated with the total available habitat. Coral-associated invertebrates do not generally use different corals indiscriminately but often exhibit speciic associations with a restricted set of different coral types (e.g., Abele & Patton 1976, Sin 1999, Stella et al. 2010). This speciicity is to be expected given the extreme diversity in gross morphology, as well as marked differences in the distribution and abundance of different coral types. However, resource specialization is a critical factor in determining a species vulnerability to disturbances, especially resource depletion (e.g., McKinney 1997). In general, those species that are more specialized in their use of 55 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES 4% 1% 1% 10% 1 taxon 2 taxa 3 taxa 4 taxa >4 taxa 84% Figure 4 The number of coral taxa used by coral-associated invertebrates based on all published records of occurrence. available resources (i.e., ecological specialists) are expected to be less able to cope with luctuations in resource availability and thus, more susceptible to extinction during major disturbances (‘the specialization-disturbance hypothesis’, vazquez & Simberloff 2002). In support of this theory, specialist species tend to predominate in less-disturbed (more stable) areas, whereas generalist species are more common in highly disturbed environments (Kitahara et al. 2000, Kassen 2002). Moreover, the responses of coral-dependent ishes to coral depletion are strongly dependent on their degree of resource specialization (Munday 2004, Pratchett et al. 2008). Among both coral-dwelling gobies (Gobiodon) and coral-feeding butterlyishes (Chaetodon), local declines in the abundance of species after extensive coral loss were directly proportional to the number of different corals that the ishes used for food or shelter (Munday 2004, Pratchett et al. 2008). The most specialized species, which tended to use only one or a few different corals, were extremely vulnerable to coral depletion and may be at risk of extinction given ongoing coral loss (Munday 2004). Establishing resource specialization for coral-associated invertebrates is often limited by significant data deiciencies. based on published records of occurrence on an identiied coral host, 84% of coral-associated invertebrates have been recorded from only one coral host taxon (Figure 4). Given the limited records of species occurrence (in some cases only one), it is not possible to assess whether these coral-associated invertebrates only utilize a single coral type or have yet to be recorded from other corals. Therefore, we can only speculate on the potential proportion of coral-associated invertebrates that are extreme specialists. If these 662 species of invertebrates are true specialists, they may face an increased risk of local and global extinction, although this also depends on the inherent vulnerability and abundance of the speciic corals used for food or shelter. Dietary specialization Coral-feeding organisms (both ishes and invertebrates) often exhibit strong selectivity for speciic coral prey, typically consuming only a small set of available coral species (reviewed by Cole et al. 2008, Rotjan & lewis 2008). Moreover, most corallivores consume corals in signiicantly different proportions to which they are available (e.g., Cox 1994, Graham 2007, Pratchett 2007). In the northern Great barrier Reef, Australia, Pratchett (2005) showed that virtually all species of coral-feeding butterlyishes (11/14 species) fed mainly on Acropora hyacinthus or Pocillopora damicornis. There was, however, marked variation in levels of specialization exhibited by different 56 CoRAl-ASSoCIATED INvERTEbRATES coral-feeding butterlyishes (Pratchett 2005). Hence, there are two important ecological questions pertaining to dietary specialization among coral-feeding organisms: (1) why are certain corals (e.g., Acropora) consistently and strongly preferred? and (2) why are some corallivores highly specialized, whereas others are comparative generalists? optimal foraging theory would predict that coral-feeding organisms would choose prey that maximize energetic return (ormond et al. 1976) and thereby selectively consume coral with the highest caloriic content (Keesing 1990), carbon-to-nitrogen ratio (Graham 2007) or high carbohydrate, protein or lipid content. Keesing (1990) explored the relationship between the nutritional value of corals and prey preferences of Acanthaster planci, and although the most highly preferred corals (e.g., Acropora spp.) were among the corals with the highest caloriic content, other nonpreferred corals from the family Faviidae also had similar nutritional value. Similar problems have hindered understanding of prey preferences among coral-feeding ishes (Graham 2007), whereby prey preferences do not consistently relate to patterns of nutritional quality. In reviewing feeding habits of Acanthaster planci, Potts (1981) suggested that coral prey consumed by A. planci may represent the least-avoided species, rather than those that are actually preferred. For example, nematocysts, mesenterial ilaments, secondary metabolites, and the antagonistic behaviour of coral symbionts may all deter corallivores from feeding on certain corals (Potts 1981). A further complication in relating feeding preferences to prey quality is that the nutritional value of corals, as measured using standard analytical techniques, may poorly relect the nutritional quality of coral to corallivores (e.g., Glynn & Krupp 1986). Glynn & Krupp (1986) conducted pairwise choice experiments to assess prey preferences of the starish Culcita novaeguineae and showed that Pocillopora meandrina was strongly preferred to Porites compressa, Montipora verrucosa and Fungia scutaria. Although the organic content of Pocillopora meandrina was the lowest of the four corals, the percent loss of organic matter after feeding by Culcita novaeguineae was the highest. This suggested that Pocillopora meandrina provides the highest energetic return for Culcita novaeguineae (Glynn & Krupp 1986), probably due to the supericial location of tissue layers and the ease with which tissues can be removed. Strong prey preferences for Acropora have also been observed for Drupella rugosa (Morton et al. 2002) and Coralliophila abbreviata (Hayes 1990) irrespective of coral community composition. Corallivorous invertebrates that are capable of feeding on a diverse range of different corals generally prefer Acroporidae (Acropora and Montipora) corals (e.g., Acanthaster planci, Pratchett 2007; Drupella cornus, Morton et al. 2002), although this may be due to conditioning, whereby species simply prefer the most abundant coral or the coral last consumed (e.g., Coralliophila abbreviata, Hayes 1990). Dietary specialization is expected to confer considerable beneits, such as increased capture and assimilation eficiency (Schoener 1971), whereby specialists are expected to outperform generalist species when preferred prey are readily accessible (e.g., Dearing et al. 2000). There was not, however, any evidence for this trade-off in the one study that speciically compared performance of specialist and generalist corallivores, based on sympatric butterlyishes in the northern Great barrier Reef (berumen & Pratchett 2008). Moreover, a number of seemingly specialist corallivores appear to be able to exploit a wide range of different corals as prey becomes scarce. Habitat specialization Coral-associated invertebrates generally exhibit strong selectivity for different coral hosts, although the range of different corals used can vary enormously. The most selective species, such as Tetralia, are known only from only one genus, Acropora (Abele & Patton 1976, Patton 1994, Sin 1999), and some tetraliids use only one or two species of Acropora, irrespective of coral abundance (Sin 1999). It would be expected that animals would select habitats that would optimize their survival and reproductive success (orians & Wittenberger 1991, Pulliam & Danielson 1991). Patterns of coral use can potentially arise from active habitat selection and differential predation risks associated 57 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES with living in different coral morphologies (lee & Sin 2009). With regard to coral morphology, tightly branching corals potentially offer greater protection from predators (vytopil & Willis 2001). Shirayama & Horikoshi (1982) recorded a greater number of free-living associates on corals with relatively ine branches and narrow interbranch space. yet even among branching corals, there is a large discrepancy in the abundance and diversity of coral associates (Stella et al. 2010). The complexity of branch growth patterns appears to be a determinant in host selection; coral with a more complex branching pattern harbours a higher diversity of associates (Stella et al. 2010). This preference for certain microhabitat characteristics results in a few coral taxa being used by a high proportion of species. Many coral associates utilize their coral host as mating sites as well, protected within coral branches (Castro 1988, Munday et al. 1997). Species of Trapezia and Tetralia crabs and Alpheus shrimps are usually found in strict mating pairs on their coral host (Castro 1978, Patton 1994, Sin 1999). Copulation in decapods usually only takes place after the female has moulted (Cheung 1968), when her soft body makes her more vulnerable to predation. Male crabs are known to guard females from predators after moulting while the female’s exoskeleton is still soft (Ryer et al. 1990, Shirley et al. 1990). once she has laid her eggs, she broods them in her abdomen until they hatch. This is a crucial time for the crabs because hatching time can range from days to weeks. Therefore, the shelter offered by coral branches can be a vital component for reproductive success and can differ in effectiveness among coral hosts. Habitat specialization has also been documented for other coral-associated invertebrates. Dai & yang (1995) documented a non-random distribution of the tube-dwelling serpulid Spirobranchus giganteus on coral reefs in southern Tawain. Four coral species, Porites lutea, P. lobata, P. lichen and Montipora informis, considered ‘competitively subordinate’ were frequently colonized by the worm, whereas most coral species were not colonized. Evidence of host selection is also apparent in molluscs. Chen et al. (2004) examined the distribution, size and reproductive characteristics of the gastropod Coralliophila violacea on two different host morphologies: branching and massive Porites. Coralliophila violacea living on branching Porites were signiicantly smaller than on massive hosts (Chen et al. 2004). Chen et al. (2004) also found that reproductive success differed because females on branching Porites had signiicantly lower fecundity than those on the massive Porites, and male-female sex changes occurred at smaller sizes on branching forms. Mokady et al. (1991) discovered that the bivalve Lithophaga lessepsiana is host speciic to Stylophora pistillata, actively choosing it over all other available corals. Stylophora pistillata was also found to trigger metamorphosis of settling larvae signiicantly more than other corals (Mokady et al. 1991). We have yet to understand the full mechanisms behind host selectivity. Further research may lead to a better understanding of how these species associations evolved. When the 869 coral-associated invertebrates in this review are grouped by the coral hosts they use, it is clear that a narrow range of coral taxa is preferred (Figure 5). of 44 coral taxa used, Pocillipora was the most preferred, with nearly 30% of species associated with this genus (Figure 5). Although this preference could be due to a bias in research on Pocillopora, resulting in more records of occurrence of the species, Pocillopora has been shown to harbour an immense diversity and abundance of associates compared with Acropora (Stella et al. 2010). The apparent high dependence of coral-associated invertebrates on certain corals could have dire consequences, depending on how those corals cope with the suite of environmental threats currently challenging coral reefs. Coral reef degradation and effects on coral-associated invertebrates Disturbance is a major determinant of the physical structure and dynamics of coral reef habitats, preventing competitive exclusion and thus maintaining high diversity (Abele 1976, Connell 1978, 58 CoRAl-ASSoCIATED INvERTEbRATES Pocillopora 300 Acropora 250 Porites Faviidae Seriatopora Fungiidae Montipora Gonipora 50 Agariciidae 100 Pavona 150 Stylophora 200 Montastrea Number of invertebrates coral-associates 350 0 Coral taxa Figure 5 Pattern of coral taxa use among coral-associated invertebrates including only those coral taxa known to be used by more than 20 species. Huston 1985, Karlson & Hurd 1993, Jones & Syms 1998, Nyström et al. 2000). Components of coral reef natural disturbance regimes include tropical cyclones, lood plumes, crown-of-thorns starish outbreaks (Pearson & Endean 1969) and the various grazing and boring activities of reef inhabitants (Hutchings 1986). Apart from the natural disturbances that reefs experience, over the past century reefs have also been subject to many anthropogenic disturbances, which are increasing in frequency and intensity. These disturbances include coastal development and sedimentation, destructive ishing practices and pollution, as well as impacts associated with climate change, such as elevated sea-surface temperatures, salinity and ocean current changes, and ocean acidiication (Goreau 1992, Sebens 1994, Wilkinson & buddemeier 1994, bryant et al. 1998, Wilkinson 1999, Jackson et al. 2001, Pandoli et al. 2003, Jones et al. 2004). These impacts have and will cause a dramatic reduction in coral cover and consequently alter the structure of populations and communities of species. Coral cover is in decline in many parts of the world, with 50% to 70% under direct threat from human activities (Goreau 1992, Hughes 1994, Sebens 1994, Wilkinson & buddemeier 1994, bryant et al. 1998, Wilkinson 1999, Gardner et al. 2003, bellwood et al. 2004), and management priorities aim to identify and remedy these threats. The impacts of reef degradation due to anthropogenic activities can be swift and devastating. land-based pollution and destructive ishing practices have caused a sharp decline of 30–60% in coral species diversity in Indonesia (Edinger et al. 1998). The loss or reduction of coral equates to a loss of vital resources to coral-associated invertebrates and other reef organisms. The subsequent effects of coral reef habitat degradation on reef organisms have been shown to be substantial. For example, Jones et al. (2004) documented serious declines in the diversity of ish assemblages due to reef degradation, with species most reliant on living coral displaying the greatest decline in abundance. As other studies have also observed the same decline of corallivorous (e.g., chaetodontids, Pratchett et al. 2006) and coral-dwelling ishes (e.g., gobies, Munday 2004), it is imperative to understand what proportion of coral-associated invertebrates rely on live corals for their long-term persistence to assess their likely sensitivity to reef degradation. The range of associations previously discussed highlights that some invertebrates associate with coral only intermittently (facultative users), while others depend on live coral for food and habitat (obligate users). of those 59 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES that have an obligate dependence on live coral, a proportion is specialized to certain coral hosts. Specialist species, which selectively utilize only one or two species of coral for food or habitat, will be particularly susceptible to reef degradation, potentially suffering greater population declines and extinction risks should their host coral decline in abundance (McKinney 1997, Pratchett et al. 2008). Therefore, the level of dependence on a coral host and how specialized an organism is to a particular coral taxon are major factors in determining the fate of coral-associated invertebrates as coral reef degradation continues. Continued coral reef degradation and the ultimate reduction in coral will probably lead to changes in the coral community composition or phase shifts, from coral-dominated to algal-dominated reefs (Hughes 1994, Hughes et al. 2007). How these changes will affect coral-associated invertebrates is not known and has only been supericially explored. Invertebrates are fairly restricted in their ability to avoid disturbance physically because most are sedentary or have limited mobility. The longterm persistence of coral-associated invertebrates will largely depend on the response diversity of different species to disturbance as well as their adaptability to a changing coral community, which may involve utilizing alternate coral hosts that are more resistant to disturbance. As biological data for the majority of coral-associated invertebrates is essentially non-existent, we have little insight regarding what the observed effects of habitat degradation might be. From the limited research into these effects, results can be conlicting. For example, Glynn et al. (1985) documented lower densities of the obligate associates Trapezia spp. on partially bleached colonies of Pocillopora damicornis, whereas Tsuchiya et al. (1992) reported an increase of the same associates after bleaching. The impact of disturbance on reef biodiversity will depend on factors such as physiology and response diversity of different species, yet ecological and biological data for most coral reef invertebrates are lacking, hampering our ability to assess which species will be most susceptible. Characteristics that some invertebrates may possess that enhance vulnerability include restricted geographic distribution or having a few small, highly fragmented populations, direct development, low fecundity, rarity and a close association with threatened taxa or threatened habitat. However, most of these factors have received little attention. because a close association with threatened taxa (i.e., coral) increases the extinction risk of species, we can make some predictions about the fate of coralassociated invertebrates with increasing reef degradation. The close association many invertebrates have with corals often relates to a certain degree of habitat specialization, which has been deemed a major factor determining the vulnerability of a species (McKinney 1997). This vulnerability is evidenced by linking four apparent recent extinctions of marine gastropods to a vulnerable habitat (Carlton 1993). Importantly, the habitat requirements of a species may differ signiicantly at various points in its life cycle; one or more life phases may be specialized for a particular habitat with limited availability. There are many examples (Pawlik 1992, Gerlach et al. 2006) of larvae that exhibit requirements for speciic types of habitat or chemical stimuli for successful settlement and even metamorphosis, such as the mussel Lithophaga lessepsiana (Mokady et al. 1991). Without speciic corals to render these cues, recruitment will probably be affected. Such inluences on recruitment will potentially affect population size, structure and persistence even if there are no obvious factors affecting the adults. It is predicted that climate change, through prolonged and intense bleaching events and subsequent habitat degradation, will cause extensive population declines and the extinction of many species, ultimately reducing biodiversity and consequently threatening the stability and resilience of coral reefs worldwide (Naeem & li 1997, Walther et al. 2002, bellwood et al. 2003, Julliard et al. 2003, bellwood et al. 2006, Przeslawski et al. 2008), but the mechanisms, whether lethal or sublethal effects, are not fully understood. Sea temperatures in many tropical regions have increased by almost 1°C over the past 100 years and are currently increasing at about 1.2°C per century (HoeghGuldberg 1999). Coral bleaching is predicted to increase in both frequency and magnitude in the years to come (Hoegh-Guldberg 1999), and rapid changes are already being seen in the community structure of coral reefs (Hughes et al. 2003). bleaching and subsequent death of corals would in 60 CoRAl-ASSoCIATED INvERTEbRATES turn result in habitat degradation and loss for many reef species, especially those considered coral specialists (Williams 1986, Kokita & Nakazono 2001, Munday 2004, Pratchett et al. 2004). Glynn et al. (1985) observed that obligate crab associates (Trapezia spp.) exhibited sublethal effects to warming that induced coral bleaching. Reproductive activity apparently declined, emigration increased, and there was a noticeable reduction in defensive behaviours. Within 2 weeks, lipid content declined by 50% in coral hosts and 78% in symbiotic crabs. bleached corals with unit crab associates may be more susceptible to disturbance and predation because these crabs have been shown to be vital to coral health through grooming and defence from predators (Glynn 1983, Stewart et al. 2006). Documented studies of coral decline have shown consequent declines in coral-associated ish assemblages and even local extinctions of coral specialists (Jones et al. 2004, Munday 2004). If ish biodiversity is affected by coral decline, and the most negative impacts are on those species deemed coral specialists, then it can be assumed that coral decline would also affect invertebrates, many of which show some degree of specialization. All coral species are not equally susceptible to coral bleaching (Coles & brown 2003). Coral colonies vary in shape and size, and these differences affect the physiological ecology of corals (Sebens 1987, Anthony et al. 2002). branching corals in particular provide a high surface area, and energy allocation to growth is disproportionately assigned to tissue growth rather than skeletal growth (Anthony et al. 2002). Fast-growing branching species (e.g., Acropora and Pocillopora) continually suffer higher bleaching mortality than slow-growing massive species (e.g., Porites and Astreopora) (brown & Suharsano 1990, Gleason 1993, Marshall & baird 2000, loya et al. 2001, Floros et al. 2004, McClanahan et al. 2004). In fact, three taxa of corals are particularly susceptible: Acropora, Pocillopora and Stylophora (loya et al. 2001, Hughes et al. 2003, McClanahan et al. 2004). Therefore, a potential outcome of increased sea-surface temperatures is a change in the relative abundance of corals. Susceptible coral taxa are often more abundant, and their loss will lead to large losses in total coral cover (Goreau et al. 2000, McClanahan et al. 2004). The apparent preference for Pocillopora may have important consequences for the many associated invertebrates. Pocillopora is known to be one of the most susceptible coral taxa to bleaching and subsequent colony mortality (brown & Suharsano 1990, Gleason 1993, Marshall & baird 2000, loya et al. 2001, Floros et al. 2004, McClanahan et al. 2004). McClanahan et al. (2004) compared bleaching susceptibility among coral taxa on Australian and Kenyan reefs using an index to indicate differences in bleaching susceptibility (score of 0–100, 0 being most resistant to bleaching and 100 being most susceptible). When this bleaching index was compared with the preferences exhibited by the coral-associated invertebrates in this review, it was clear Pocillopora is the coral taxon used by the highest proportion of coral associates and is one of the most susceptible coral taxa to bleaching (Figure 6). Moreover, it was found that invertebrates exhibit a heavy reliance on other susceptible coral taxa as well. of all the coral-associated invertebrates found in this review, 53% (462 species) utilize three branching coral species considered to be the most susceptible coral taxa: Acropora, Pocillopora, and Stylophora (Figure 6). Therefore, the decline of even three genera of coral, with prolonged and frequent bleaching, could potentially result in an immense loss of reef biodiversity. The widely predicted scenario in which coral reefs become dominated by algae will result in systems that are unlikely to host the highly specialized invertebrate associates intimately associated with corals. Further threats to coral-associated invertebrates Coral-associated invertebrates can and will be directly affected by anthropogenic and environmental changes irrespective of the impacts to their coral host. Many coral-reef invertebrate species are commercially and recreationally harvested for food, bait and ornamentation. These species include molluscs such as oysters, scallops, squid, octopus and abalone; crustaceans such as rock lobsters and crabs; and holothurian echinoderms (trepang or bêche de mer) (Australian bureau of Agricultural 61 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES 275 Pocillopora 250 Number of species that use coral 225 200 175 150 Acropora 125 100 75 Stylophora Faviidae 50 Fungiidae Porites Goniopora Porites Pavona Montipora Astreopora Goniastrea Galaxea 25 0 0 10 20 30 40 50 60 70 80 90 100 Australian bleaching index Figure 6 Scatterplots of the Australian bleaching Index for coral taxa and the number of invertebrate species dependent on coral taxa. (Australian bleaching Index taken from McClanahan et al. 2004 and; species numbers taken from collated published data.) and Resource Economics [AbARE] 1998). bait harvests typically include worms, intertidal molluscs, crustaceans and ascidians. Species collected for jewellery include shelled molluscs, such as trochus, cowry and cone shells, as well as coral, some echinoderms and crustaceans. Although this trade is strictly controlled in parts of the world, such as Australia, many developing countries have few, if any, controls. Moreover, many other species are collected live for the aquarium trade. Direct exploitation of marine invertebrates for human consumption is a valuable industry, worth more than A$1.3 billion in Australia alone (AbARE 1998) and in many developing countries would be much higher as part of subsistence ishing. Due to the small size of most coral associates, they are not as highly valued as food items but are still collected as bait, ornamentation and the live aquarium trade. However, overexploitation is not considered an immediate threat (Hutchings et al. 2007). Marine invertebrate biodiversity also represents a vast resource of novel bioactive compounds, such as the anticancer agent developed from the Caribbean sponge Cryptotheca crypta (McConnell et al. 1994) and the potential use of the toxin of the cone snail to treat neuropsychiatric disorders such as Alzheimer’s and Parkinson’s diseases (olivera et al. 1990). However, as the progress of drug discovery in the marine environment is slow, overharvesting for medicinal purposes is not a threat, and many would argue that the potential beneits could far outweigh the risks. Climate change will subject coral reefs to a suite of environmental changes, including changes in ocean currents, a rise in the sea-surface temperature, changes in ocean chemistry, increased rainfall and freshwater plumes, a rising sea level and increased irradiance (Hoegh-Guldberg 1999, Hughes et al. 2003, West & Salm 2003, Munday et al. 2007). The impacts of climate change on coral reef invertebrates are almost entirely unknown (but see Przeslawski et al. 2008). Rising sea level has the potential to affect the distribution of intertidal invertebrates, allowing them to expand landwards, providing that suitable habitat is available. Changes in ocean currents may have serious implications for marine invertebrate populations. Many marine invertebrates have a pelagic larvae stage that may extend from a few hours to many weeks (levin 2006). The strength and direction 62 CoRAl-ASSoCIATED INvERTEbRATES of ocean currents is critical for larvae to return to either their natal reef or a nearby suitable reef. Changing the direction and strength of ocean currents may carry larvae to unsuitable habitats where survival will be unlikely. Increases in storm activity and associated changes in salinity will affect both larval survival and the invertebrates living in shallow waters, including lagoonal reefs. Temperature is considered to be the most inluential factor in the physiological processes of marine animals and thus greatly shapes their biogeographic distributions (Clarke 2003, Mueter & litzow 2008, Richardson 2008, Tewksbury et al. 2008). Temperature is a critical factor for invertebrates and determines growth and reproduction rates and survival. The developmental and growth rates of marine invertebrates show a strong positive correlation with temperature (Fujisawa & Shigei 1990, Palmer 1994, Reitzel et al. 2004, o’Connor et al. 2007, Sheppard brennand et al. 2010) until the thermal threshold is reached. Therefore, temperature increases associated with climate change may at irst accelerate the growth rates of many species living in the middle of their temperature tolerance range, whereas those living closer to their thermal maxima may become extinct (Precht & Aronson 2004, Greenstein & Pandoli 2008). For many species, it is the climate extremes that are critical, and as lough & barnes (1990) indicated, it is the extremes that are projected to increase signiicantly. Tropical species typically have a lower tolerance to temperature variation than temperate species (Compton et al. 2007) because they may already live near their thermal optima (Tewksbury et al. 2008). Furthermore, symbioses in tropical species may be less stable during thermal luctuations than those in temperate species (Muller-Parker & Simon 2001). Invertebrates, particularly crustaceans, are sensitive to salinity and thermal changes. Abele (1976, 1979) found that the crustacean associates of Pocillopora were generally more sensitive to environmental extremes, particularly salinity changes, than their coral hosts. Increased temperatures associated with El Niño were found to have signiicant negative effects on crustacean associates, causing a massive decline in abundance (Glynn & D’Croz 1990). upwellings and oxygen depletion can kill crab and shrimp associates but may only cause partial mortality to the corals (Abele 1976, 1979, Glynn et al. 1985). Echinoderms are also sensitive to temperature increases. Complete developmental failure was observed among sea urchins in manipulated high temperatures (4–6°C above normal) (byrne et al. 2009). An increase in ocean temperature therefore would have deleterious effects on the development, growth and reproduction of marine invertebrates. ocean acidiication is also a direct threat to marine invertebrates, particularly species with calcareous shells (i.e., molluscs, echinoderms, crustaceans, bryozoans, serpulid polychaetes, foraminiferans, and sponges as well as corals) especially if ocean pH falls below 7.5 (Raven et al. 2005, Kleypas et al. 2006, Gazeau et al. 2007, but see Wood et al. 2008). vulnerability to ocean acidiication on larval development and calciication is highly variable among crustaceans and molluscs (as reviewed by byrne 2011). Crustaceans may be more resilient to ocean acidiication due to the high organic content of chitin in their shells (Derry & Arnott 2007), as opposed to the more susceptible aragonite skeleton of molluscs (byrne 2011). bivalve molluscs appear to be particularly susceptible to decreases in ocean pH. High levels of atmospheric carbon dioxide have detrimental effects on the shell synthesis of larval bivalves (Kurihara et al. 2008). Adults have also been shown to suffer deleterious effects. Carbon dioxide levels equivalent to 740 ppm caused the calciication rate in the mussel Mytilus edulis to decrease by 25% (Gazeau et al. 2007). Echinoderms are also under direct threat of ocean acidiication. Sea urchins have exhibited a marked decrease in fertilization success, developmental rates, and larval size with increasing carbon dioxide concentrations (Kurihara & Shirayama 2004, Sheppard brennand et al. 2010). The direct threats climate change poses to marine invertebrates may be severe, and it is important to consider that many threats will be acting synergistically and with apparent selectivity on certain invertebrate groups (reviewed by byrne 2011). The onslaught of temperature increases, reduced salinity and ocean acidiication as well as the consequences of habitat degradation will be trying, at best, on the persistence of coralassociated invertebrate populations. 63 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Conclusions and further directions This review indicates that coral-associated invertebrates are a diverse group of reef organisms that are of concern as a consequence of declining reef health. Although most invertebrate groups are poorly described and in urgent need of taxonomic studies, a number of important generalizations have begun to emerge. The biodiversity of coral-associated invertebrates is dominated by the great number of arthropods, particularly the decapod crustaceans (Abele 1974, Patton 1974, Abele 1976, Abele & Patton 1976, Austin et al. 1980, Stella et al. 2010). Approximately half of all known coral-associated invertebrates have an obligate association with coral hosts. The data available also indicate that the majority of many coral-associated invertebrates exhibit a high degree of habitat specialization, living in symbiosis with particular corals or utilizing a subset of coral prey. branching corals of the genera Acropora, Pocillopora and Stylophora harbour the highest diversity of associates. Importantly, these coral taxa are also among the most susceptible to bleaching. Climate change-induced coral bleaching will lead to a reduction or a change in the distribution and abundance of corals, with a greater impact on corals most susceptible to bleaching. Therefore, it can be hypothesized that declines in coral host abundance will lead to declines in abundance diversity of coral-associated invertebrates. Although many coral-associated invertebrates depend on corals, it is also apparent that some coral-associated invertebrates are in turn fundamental to the itness and persistence of their host corals. Knowledge of interactions and feedbacks between corals and coral-associated invertebrates is currently limited to a few speciic examples (e.g., Pratchett 2001, Stewart et al. 2006), but there is the potential that many coral-associated species, even seemingly deleterious species, could have beneicial effects for corals (e.g., Mokady et al. 1998). Invertebrates may perform many important functional roles on coral reefs, with the potential to inluence coral health and serve as an important trophic link between corals and other reef organisms. Scientiic monitoring of ecosystem health on coral reefs needs to encompass the vast diversity and abundance of coral reef invertebrates, recognizing their functional roles and susceptibility to climate change. We are currently faced with a critical lack of knowledge, which limits our ability to protect and continue to beneit from coral reef biodiversity. Success will depend on effective research and management of coral reefs that includes all components vital to ecosystem function. It is unfortunate that coral-associated invertebrates exhibit many traits that enhance their risk of extinction via direct and indirect threats due to climate change and subsequent habitat degradation. The conclusion that coral reef biodiversity as a whole is under a severe threat is inescapable. The challenge will be to increase our focus on coral-associated invertebrate communities, the largest component of coral reef biodiversity, and fully assess their role in coral reef ecosystem function and resilience. Acknowledgements We thank the Australian government, Department of Climate Change, the Australian Institute of Marine Science, the linnean Society of New South Wales, the School of Marine and Tropical biology at James Cook university and the ARC Centre of Excellence for Coral Reef Studies for their support of this study. Thanks to R. Gibson for comments on the manuscript. References Abele, l.G. 1974. Species diversity of decapod crustaceans in marine habitats. Ecology 55, 156–161. Abele, l.G. 1976. Comparative species richness in luctuating and constant environments: coral associated decapod crustaceans. Science 192, 461–463. 64 CoRAl-ASSoCIATED INvERTEbRATES Abele, l.G. 1979. The community structure of coral-associated decapod crustaceans in variable environments. In Ecological Processes in Coastal and Marine Systems, R.J. livingston (ed.). New york: Plenum Press, 265–287. Abele, l.G. & Patton, W.K. 1976. The size of coral heads and community biology of associated decapod crustaceans. Journal of Biogeography 3, 35–47. Achituv, y. 2001. Cantellius alphonei n.sp., a new coral-inhabiting barnacle (Cirripedia, Pyrgomatidae) of Montipora. Crustaceana 74, 617–626. Achituv, y. & Hoeksema, b.W. 2003. Cantellius cardenae spec. nov. (Cirripedia: Pyrgomatinae) from Acropora (Isopora) brueggemanni (brook, 1893) (Anthozoa: Acroporidae), a case of host speciicity in a generalist genus. Zoologische Mededelingen 77, 1–14. Achituv, y. & Newman, W.A. 2002. The barnacles of Astreopora (Cirripedia, Pyrgomatini/Scleractinia, Acroporidae): organization plans, host speciicity, species-richness and geographic range. Journal of Natural History 36, 391–406. Achituv, y., Tsang, l.M. & Chan, b.K.K. 2009. A new species of Cantellius and a redescription of C. sextus (Hiro, 1938) (Cirripedia, balanomorpha Pyrgomatidae) from the elephant skin coral, Pachyseris speciosa (Dana, 1846) (Scleractinia, Agariciidae) from Taiwan. Zootaxa 2022, 15–28. Aeby, G.S. & Santavy, D.l. 2006. Factors affecting susceptibility of the coral Montastraea faveolata to blackband disease. Marine Ecology Progress Series 318, 103–110. Aerts, l.A.M & van Soest, R.W.M 1997. Quantiication of sponge/coral interactions in a physically stressed reef community, NE Colombia. Marine Ecology Progress Series 148, 125–134. Anker, A., Ahyong, S.T., Noël, P. & Palmer, A.R. 2006. Morphological phylogeny of alpheid shrimps: parallel preadaptation and the origin of a key morphological innovation, the snapping claw. Evolution 60, 2507–2528. Anthony, K.R.N., Connolly, S.R. & Willis, b.l. 2002. Comparative analysis of energy allocation to tissue and skeletal growth in corals. Limnology and Oceanography 47, 1417–1429. Antonius, A. & Riegl, b. 1997a. A possible link between coral diseases and a corallivorous snail (Drupella cornus) outbreak in the Red Sea. Atoll Research Bulletin 447, 1–9. Antonius, A. & Riegl, b. 1997b. Coral diseases and Drupella cornus invasion in the Red Sea. Coral Reefs 17, 48. Austin, A.D., Austin, S.A. & Sale, P.F. 1980. Community structure of the fauna associated with the coral Pocillopora damicornis (l.) on the Great barrier Reef. Australian Journal of Marine and Freshwater Research 31, 163–174. Australian bureau of Agricultural and Resource Economics (AbARE). 1998. Australian isheries statistics 1998. Canberra: Australian bureau of Agricultural and Resource Economics. bak, R.P.M. 1990. Patterns of echinoid bioerosion in two Paciic coral reef lagoons. Marine Ecology Progress Series 66, 267–272. barneah, o., brickner, I., Hooge, M., Weis, v.M., laJeunesse, T.C. & benayahu, y. 2007. Three party symbiosis: acoelomorph worms, corals and unicellular algal symbionts in Eilat (Red Sea). Marine Biology 151, 1215–1223. bartolomaeus, T. & balzer, I. 1997. Convolutriloba longiissura, sp. nov. (Acoela)—irst case of longitudinal ission in Platyhelminthes. Microfauna Marina 11, 7–18. baums, I.b., Miller, M.W. & Szmant, A.M. 2003a. Ecology of a corallivorous gastropod on two scleractinian hosts. I: Population structure of snails and corals. Marine Biology 142, 1083–1091. baums, I.b., Miller, M.W. & Szmant, A.M. 2003b. Ecology of a corallivorous gastropod on two scleractinian hosts. II: Feeding, respiration and growth. Marine Biology 142, 1093–1101. bautista-Guerrero, E., Carballo, J.l., Cruz-barraza, J.A. & Nava, H.H. 2006. New coral reef boring sponges (Hadromerida: Clionaidae) from the Mexican Paciic ocean. Journal of the Marine Biological Association of the United Kingdom 86, 963–970. bellwood, D.R., Hoey, A.S., Ackerman, J.l. & Depczynski, M. 2006. Coral bleaching, reef ish community phase shifts and the resilience of coral reefs. Global Change Biology 12, 1587–1594. bellwood, D.R., Hoey, A.S. & Choat, J.H. 2003. limited functional redundancy in high diversity systems; resilience and ecosystem function in coral reefs. Ecology Letters 6, 281–285. bellwood, D.R., Hughes, T.P., Folke, C. & Nyström M. 2004. Confronting the coral reef crisis. Nature 429, 827–833. 65 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES bergsma, G.S. 2009. Tube-dwelling coral symbionts induce signiicant morphological change in Montipora. Symbiosis 49, 143–150. berumen, M.l. & Pratchett, M.S. 2008. Trade-offs associated with dietary specialisation in corallivorous butterlyishes (Chaetodontidae: Chaetodon). Behavioural Ecology and Sociobiology 62, 989–994. birkeland, C. 1982. Terrestrial runoff as a cause of outbreaks of Acanthaster planci Echinodermata: Asteroidea). Marine Biology 69, 175–185. birkeland, C. 1989. The inluence of echinoderms on coral reef communities. In Echinoderm Studies 3, M. Jangoux & J.M. lawrence (eds). Rotterdam: balkema, 1–79. birkeland, C. & lucas, J.S. 1990. Acanthaster planci: Major Management Problem of Coral Reefs. boston: CRC Press. black, R. & Prince, J. 1983. Fauna associated with the coral Pocillopora damicornis at the southern limit of its distribution in Western Australia. Journal of Biogeography 10, 135–152. boucher, l.M. 1986. Coral predation by muricid gastropods of the genus Drupella at Enewetak, Marshall Islands. Bulletin of Marine Science 38, 9–11. bouchet, P. 2006. The magnitude of marine biodiversity. In The Exploration of Marine Biodiversity Scientiic and Technological Challenges, C.M. Duarte (ed.). bilbao, Spain: Fundación bbvA, 31–62. bouchet, P., lozouet, P., Maestrati, P. & Heros, v. 2002. Assessing the magnitude of species richness in tropical marine environments: exceptionally high numbers of molluscs at a New Caledonia site. Biological Journal of the Linnean Society 75, 421–436. bowers, R.l. 1970. The behavioral ecology of Alpheus clypeatus Coutiere (Decapoda, Alpheidea). Dissertation Abstracts 31, 50–70. branham, J.M., Reed, S.A. & bailey, J.H. 1971. Coral-eating sea stars Acanthaster planci in Hawaii. Science 172, 1155–1157. brauer, R.W., Jordan, M.R. & barnes, D.J. 1970. Triggering of the stomach eversion relex of Acanthaster planci by coral extracts. Nature 228, 344–346. brawley, S.H. & Adey, W.H. 1982. Coralliophila abbreviata: a signiicant corallivore! Bulletin of Marine Science 32, 595–599. brickner, I., Simon-blecher, N. & Achituv, y. 2010. Darwin’s Pyrgoma (Cirripedia) revisited: revision of the Savignium group, molecular analysis and description of new species. Journal of Crustacean Biology 30, 266–291. britayev, T.A. 1981. Two new species of commensal polynoids (Polychaeta: Polynoidae) and bibliography on polychaetes, symbionts of Coelenterata. Zoologicheski Zhurnal 60, 817–824. brodie, J., Fabricius, K., De’ath, G. & okaji, K. 2005. Are increased nutrient inputs responsible for more outbreaks of crown-of-thorns starish? An appraisal of the evidence. Marine Pollution Bulletin 51, 266–278. brown, b.E. & Suharsono 1990. Damage and recovery of coral reefs affected by El Niño related seawater warming in the Thousand Islands, Indonesia. Coral Reefs 8, 163–170. bruce, A.J. 1966. Notes on some Indo-Paciic Pontoniinae, XI: a re-examination of Philarius lophos barnard, with the designation of a new genus, Ischnopontonia. Bulletin of Marine Science 16, 584–598. bruce, A.J. 1969. Preliminary descriptions of sixteen new species of the genus Periclimenes Costa 1844 (Crustacea, Decapoda Natantia, Pontoniinae). Zoologische Mededelingen Uitgegeven door het Rijksmuseum van Natuurlijke Historie te Leiden 43, 253–278. bruce, A.J. 1972a. on the association of the shrimp Racilius compressus Paulson (Decapoda, Alpheidae) with the coral Galaxea clavus (Dana). Crustaceana 22, 92–93. bruce, A.J. 1972b. A report on a small collection of pontoniid shrimps from Fiji, with the description of a new species of Coralliocaris Stimpson (Crustacea, Decapoda, Natantia, Pontoniinae). Paciic Science 26, 63–86. bruce, A.J. 1973. Notes on some Indo-Paciic Pontoniinae, XXIII: Tectopontonia maziwiae gen. nov., sp. nov., a new coral associate from Tanganyika (Decapoda, Palaemonidae). Crustaceana 24, 169–180. bruce, A.J. 1974a. Coralliocaris viridis sp. nov., a preliminary note (Decapoda Natantia. Pontoniinae). Crustaceana 26, 222 only. bruce, A.J. 1974b. A report on a small collection of pontoniinid shrimps from the Island of Farquhar (Decapoda, Palaemonidae). Crustaceana 205, 189–203. bruce, A.J. 1974c. observations upon some specimens of the genus Periclimenaeus borradaile (Decapoda, Natantia, Pontoniinae) originally described by G. Nobili. Bulletin du Museum National d’Histoire Naturelle, Series 3, 258 (Zoologie 180), 1557–1583. 66 CoRAl-ASSoCIATED INvERTEbRATES bruce, A.J. 1974d. A synopsis of the pontoniid shrimp fauna of Central East Africa. Journal of the Marine Biological Association of India 16, 462–490. bruce, A.J. 1976. A report on a small collection of shrimps from the Kenyan National Marine Parks at Malindi, with notes on selected species. Zoologische Verhandelingen Leiden 145, 1–72. bruce, A.J. 1977. The hosts of the coral-associated Indo-West-Paciic pontoniine shrimps. Atoll Research Bulletin 205, 1–19. bruce, A.J. 1978. The re-examination of some pontoniine shrimp types irst described by l.A. borradaile (Decapoda, Palaemonidae). Crustaceana 34, 251–268. bruce, A.J. 1979. Notes on some Indo-Paciic Pontoniinae, XXXI Periclimenes magniicus sp. nov., a coelenterate associate from the Capricorn Islands (Decapoda, Palaemonidae). Crustaceana Supplement 5, 195–208. bruce, A.J. 1984. Marine caridean shrimps of the Seychelles. In Biogeography and Ecology of the Seychelles Islands, D.R. Stoddart (ed.). The Netherlands: The Hague, 141–170. bruce, A.J. 1997. A new pontoniine shrimp genus (Crustacea: Decapoda) from the yemen, with a note on other species. Journal of Natural History 31, 1213–1222. bruce, A.J. 2005. Pontoniine shrimps from Papua New Guinea, with description of two new genera, Cainonia and Colemonia (Crustacea: Decapoda: Palaemonidae). Memoirs of the Queensland Museum 51, 333–383. bruno, J.F. & Selig, E.R. 2007. Regional decline of coral cover in the Indo-Paciic: timing, extent, and subregional comparisons. PLoS ONE 2, e711. bryant, D., burke, l., McManus, J. & Spalding, M. 1998. Reefs at Risk: A Map-Based Indicator of Threats to the World’s Coral Reefs. Washington, DC: World Resources Institute. byrne, M. 2011. Impact of ocean warming and ocean acidiication on marine invertebrate life history stages: vulnerabilities and potential for persistence in a changing ocean. Oceanography and Marine Biology An Annual Review 49, 1–42. byrne, M., Ho, M., Selvakumaraswamy, P., Nguyen, H., Dworjanyn, S., & Davis, A. 2009. Temperature, but not pH, compromises sea urchin fertilization and early development under near-future climate change scenarios. Proceedings of the Royal Society London Series B 276, 1883–1888. Carballo, J.l., Hepburn, l., Nava, H. & Cruz-barraza, J.A. 2007. Coral reefs boring Aka species (Porifera: Phloeodictyidae) from Mexico with description of Aka cryptica sp. nov. Journal of the Marine Biological Association of the United Kingdom 87, 1477–1484. Carlton, J.T. 1993. Neoextinctions of marine invertebrates. American Zoologist 33, 499–509. Carpenter, R.C. 1997. Invertebrate predators and grazers. In The Life and Death of Coral Reefs, C. birkeland (ed.). New york: Chapman and Hall, 198–248. Castro, P. 1976. brachyuran crabs symbiotic with scleractinian corals; a review of their biology. Micronesica 12, 99–110. Castro, P. 1978. Movements between coral colonies in Trapezia ferruginea (Crustacea: brachyura), an obligate symbiont of scleractinian corals. Marine Biology 46, 237–245. Castro, P. 1988. Animal symbioses in coral reef communities: a review. Symbiosis 5, 161–184. Castro, P., Ng, P.K.l. & Ahyong, S.T. 2004. Phylogeny and systematics of the Trapeziidae Miers, 1886 (Crustacea: brachyura), with the description of a new family. Zootaxa 643, 1–70. Chang, K., Chen, y. & Chen, C. 1987. Xanthid crabs in the corals Pocillopora damicornis and P. verrucosa of Southern Taiwan. Bulletin of Marine Science 41, 214–220. Chen, M., Soong, K. & Tsai, M. 2004. Host effect on size structure and timing of sex change in the coralinhabiting snail Coralliophila violacea. Marine Biology 144, 287–293. Cheng, y.R. & Dai, C.F. 2010. Endosymbiotic copepods may feed on zooxanthellae from their coral host, Pocillopora damicornis. Coral Reefs 29, 13–18. Cheng, y.R., Ho, J.S. & Dai, C.F. 2007. Two new species of Xariia Humes, 1960 (Copepoda, Xariidae) associated with corals in Taiwan. Crustaceana 80, 135–1144. Cheng, y.R., Ho, J.S. & Dai, C.F. 2009. Orstomella yaliuensis n. sp., a xariiid copepod (Crustacea) parasitic in the polyps of hump coral Porites lutea Milne Edwards & Haime off Taiwan. Systematic Parasitology 74, 17–21. Chesher, R.H. 1969. Destruction of Paciic corals by the sea star Acanthaster planci. Science 18, 280–283. Cheung, T.S. 1968. Trans-molt retention of sperm in the female stone crab, Menippe mercenaria (Say). Crustaceana 15, 117–120. 67 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Choat, J.H. & bellwood, D.R. 1991. Reef ishes: their history and evolution. In The Ecology of Fishes on Coral Reefs, P.F. Sale (ed.). San Diego, California: Academic Press, 39–68. Clarke, A. 2003. Costs and consequences of evolutionary temperature adaptation. Trends in Ecology and Evolution 18, 573–581. Coker, D.J., Pratchett, M.S. & Munday, P.l. 2009. Coral bleaching and habitat degradation increase susceptibility to predation for coral-dwelling ishes. Behavioral Ecology 20, 1204–1210. Cole, A.J., lawton, R.J., Pratchett, M.S. & Wilson, S.K. 2010. Chronic coral consumption by butterlyishes. Coral Reefs doi:10.1007/s00338-010-0674-6. Cole, A.J., Pratchett, M.S. & Jones, G.P. 2008. Diversity and functional importance of coral-feeding ishes on tropical coral reefs. Fish and Fisheries 9, 286–307. Cole, A.J., Pratchett, M.S. & Jones, G.P. 2009. Coral-feeding wrasse scars massive Porites colonies. Coral Reefs 28, 207 only. Coles, S.l. 1980. Species diversity of decapods associated with living and dead reef coral Pocillopora meandrina. Marine Ecology Progress Series 2, 281–291. Coles, S.l. & brown, b.E. 2003. Coral bleaching-capacity for acclimatization and adaptation. Advances in Marine Biology 46, 183–223. Coleman, N. 2003. 2002 Sea Shells. Springwood, Queensland, Australia: Neville Coleman’s underwater Geographic. Colgan, M.W. 1985. Growth rate reduction and modiication of a coral colony by a vermetid mollusc, Dendropoma maxima. Proceedings of the 5th International Coral Reef Symposium 6, 205–210. Colgan, M.W. 1987. Coral reef recovery on Guam (Micronesia) after catastrophic predation by Acanthaster planci. Ecology 68, 1592–1605. Collins, A.R.S. 1975. biochemical investigation of two responses involved in the feeding behaviour of Acanthaster planci (l.). III. Food preferences. Journal of Experimental Marine Biology and Ecology 17, 87–94. Compton, T.J., Rijkenberg, M.J.A., Drent, J. & Persma, T. 2007. Thermal tolerance ranges and climate variability: a comparison between bivalves from differing climates. Journal of Experimental Marine Biology and Ecology 352, 200–211. Connell, J.H. 1978. Diversity in tropical rain forests and coral reefs. Science 199, 1302–1310. Cox, E.F. 1994. Resource use by corallivorous butterlyishes (Family Chaetodontidae) in Hawaii. Bulletin of Marine Science 54, 535–545. Cruz-barraza, J.A. & Carballo, J.l. 2008. Taxonomy of sponges (Porifera) associated with corals from the Mexican Paciic ocean. Zoological Studies 47, 741–758. Dai, C. & yang, H. 1995. Distribution of Spirobranchus giganteus corniculatus (Hove) on the coral reefs of southern Taiwan. Zoological Studies 34, 117–125. Dalton, S.J. & Godwin, S. 2006. Progressive coral tissue mortality following predation by a coralivorous nudibranch (Phestilla sp.). Coral Reefs 25, 529 only. Dana, T. & Wolfson, A. 1970. Eastern Paciic crown-of-thorns starish populations in the lower Gulf of California. Transactions of the San Diego Society of Natural History 16, 83–90. De Grave, S. 2000. Caridean shrimps (Crustacea, Decopoda) from Hansa bay, Papua New Guinea: Palaemonidae and Gnathophyllidae. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique Biologie 70, 119–148. De’ath, G. & Moran, P.J. 1998. Factors affecting the behaviour of crown-of-thorns starish (Acanthaster planci l.) on the Great barrier Reef. 2. Feeding preferences. Journal of Experimental Marine Biology and Ecology 220, 107–126. Dearing, M.D., Mangione, A.M. & Karasov, W.H. 2000. Diet breadth of mammalian herbivores: nutrient versus detoxiication constraints. Oecologia 123, 397–405. Debelius, H. 2001. Crustacea Guide of the World. Frankfurt, Germany: IKAN-unterwasserarchiv. Delbeek, J.C. & Sprung, J. 1997. The Reef Aquarium: A Comprehensive Guide to the Identiication and Care of Tropical Marine Invertebrates. Coconut Grove, Florida: Ricordea. Derry, A.M. & Arnott, S.E. 2007. Adaptive reversals in acid tolerance in copepods from lakes recovering from historical stress. Ecological Applications 17, 1116–1126. Devantier, l.M. & Endean, R. 1988. The scallop Pedum spondyloideum mitigates the effects of Acanthaster planci predation on the host coral Porites: host defence facilitated by exaptation? Marine Ecology Progress Series 47, 293–301. 68 CoRAl-ASSoCIATED INvERTEbRATES Devantier, M., Reichelt, R.E. & bradbury, R.H. 1986. Does Spirobranchus giganteus protect host Porites from predation by Acanthaster planci: predator pressure as a mechanism of coevolution? Marine Ecology Progress Series 32, 307–310. Dilly, P.N. & Ryland, J.S. 1985. An intertidal Rhabdopleura (Hemichordata, Pterobranchia) from Fiji. Journal of Zoology, London 205, 611–623. Dojiri, M. 1988. Isomolgus desmotes, new genus, new species (lichomolgidae), a gallicolous poecilostome copepod from the scleractinian coral Seriatopora hystrix Dana in Indonesia, with a review of gallinhabiting crustaceans of Anthozoans. Journal of Crustacean Biology 8, 99–109. Duffy, J.E. 2003. biodiversity loss, trophic skew and ecosystem functioning. Ecology Letters 6, 680–687. Dulvy, N.K., Ellis, J.R., Goodwin, N.b., Grant, A., Reynolds, J.D. & Jennings, S. 2004. Methods of assessing extinction risk in marine ishes. Fish and Fisheries 5, 255–275. Earle, S.A. 1991. Sharks, squids, and horseshoe crabs-the signiicance of marine biodiversity. Bioscience 47, 506–509. Ebbs, N.K. 1966. The coral-inhabiting polychaetes of the northern Florida reef tract. Part I. Aphroditidae, Polynoidae, Amphinomidae, Eunicidae, and lysaretidae. Bulletin of Marine Science 16, 485–555. Edinger, E.N., Jompa, J., limmon, G.v., Widjatmoko, W. & Risk, M. 1998. Reef degradation and coral biodiversity in Indonesia: effects of land-based pollution, destructive ishing practices and changes over time. Marine Pollution Bulletin 36, 617–630. Edmondson, C.H. 1933. Cryptochirus of the central Paciic. Occasional Papers of the Bernice P. Bishop Museum 10, 1–23. Edwards, A. & Emberton, H. 1980. Crustacea associated with the scleractinian coral Stylophora pistillata (Esper), in the Sudanese Red Sea. Journal of Experimental Marine Biology and Ecology 42, 225–240. Fabricius, K.E. 2005. Effects of terrestrial runoff on the ecology of corals and coral reefs: review and synthesis. Marine Pollution Bulletin 50, 125–146. Fabricius, K.E., okaji, K. & De’ath, A.G. 2010. Three lines of evidence to link outbreaks of the crown-ofthorns seastar Acanthaster planci to the release of larval food limitation. Coral Reefs 29, 593–605. Fang, l.S. & Shen, P. 1988. A living mechanical ile: the burrowing mechanism of the coral-boring bivalve Lithophaga nigra. Marine Biology 97, 349–354. Fauchald, K. 1992. A review of the genus Eunice (Eunicidae: Polychaeta) based upon type material. Smithsonian Contributions to Zoology 523, 1–422. Fize, A. & Serène, R. 1956. Note préliminaire sur huit éspèces nouvelles, dont une d’un genre nouveau, d’hapalocarcinides. Bulletin de la Société Zoologique de France 80, 375–378. Fize, A. & Serène, R. 1957. les hapalocarcinides du vietNam. Archives du Museum National d’Histoire Naturelle Paris 7, 1–202. Floros, C.D., Samways, M.J. & Armstrong, b. 2004. Taxonomic patterns of bleaching within a South African coral assemblage. Biodiversity and Conservation 13, 1175–1194. Fonseca, A.C. & Cortés, J. 1998. Coral borers of the Eastern Paciic: Aspidosiphon (A.) elegans (Sipuncula: Aspidosiphonidae) and Pomatogebia rugosa (Crustacea: upogebiidae). Paciic Science 52, 170–175. Fossa, S.v. & Nilsen, A.J. 2000. The Modern Reef Aquarium. bornheim, Germany: birgit Schmettkamp verlag. Fujino, T. & Miyake, S. 1972. A new pontoniinid shrimp of the genus Coralliocaris Stimpson from Taiwan (Crustacea, Decapoda, Pontoniinae). Occasional Papers of the Zoological Laboratory, Faculty of Agriculture Kyushu University 3, 91–98. Fujisawa, H. & Shigei, M. 1990. Correlation of embryonic temperature sensitivity of sea urchins with spawning season. Journal of Experimental Marine Biology and Ecology 136, 123–139. Galil, b.S. & Clark, P.F. 1988. on a collection of Acropora-inhabiting trapeziids (Crustacea brachyura Xanthoidea) from East Africa. Tropical Zoology 1, 137–151. Galil, b. & Takeda, M. 1986. Resurrection of the genus Jonesius and the establishment of a new genus: commensal crabs associated with corals from the Indo-Paciic ocean. Bulletin of the National Science Museum Tokyo 12, 163–171. Galil, b.S. & vannini, M. 1990. Research on the coast of Somalia. Xanthidae, Trapeziidae, Carpiliidae, Menippidae (Crustacea brachyura). Tropical Zoology 3, 21–56. Gardner, T.A., Côté, I.M., Gill, J.A., Grant, A. & Watkinson, A.R. 2003. long-term region-wide declines in Caribbean corals. Science 301, 958–960. Garpe, K.C., yahya, S.A.S., lindahl, u. & Öhman, M.C. 2006. long-term effects of the 1998 coral bleaching event on reef ish assemblages. Marine Ecology Progress Series 315, 237–247. 69 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Gaston, K. 1991. The magnitude of global insect species richness. Conservation Biology 5, 283–296. Gazeau, F., Quiblier, C., Jansen, J.M., Gattuso, J.P., Middelburg, J.J. & Heip, C.H.R. 2007. Impact of elevated Co2 on shellish calciication. Geophysical Research Letters 34, l07603. doi:10.1029/2006Gl028554. Gerlach, G., Atema, J., Kingsford, M.J., black, K.P. & Millers-Sims, v. 2006. Smelling home can prevent dispersal of reef ish larvae. Proceedings of the National Academy of Sciences of the United States of America 104, 858–863. Gittenberger, A. 2003. The wentletrap Epitonium hartogi spec. nov. (Gastropoda: Epitoniidae), associated with bubble coral species, Plerogyra spec. (Scleractinia: Euphylliidae), off Indonesia and Thailand. Zoologische Verhandelingen Leiden 345, 139–150. Gittenberger, A. & Gittenberger, E. 2005. A hitherto unnoticed adaptive radiation: epitoniid species (Gastropoda: Epitoniidae) associated with corals (Scleractinia). Contributions to Zoology 74, 125–203. Gittenberger, A., Goud, J. & Gittenberger, E. 2000. Epitonium (Gastropoda: Epitoniidae) associated with mushroom corals (Scleractinia: Fungiidae) from Sulawesi, Indonesia, with the description of four new species. The Nautilus 114, 1–13. Gleason, M.G. 1993. Effects of disturbance on coral communities: bleaching in Moorea, French Polynesia. Coral Reefs 12, 193–201. Glynn, P.W. 1973. Acanthaster: effect on coral reef growth in Panamá. Science 180, 504–506. Glynn, P.W. 1974. The impact of Acanthaster on corals and coral reefs in the eastern Paciic. Environmental Conservation 1, 295–303. Glynn, P.W. 1976. Some physical and biological determinants of coral community structure in the Eastern Paciic. Ecological Monographs 46, 431–456. Glynn, P.W. 1982. Acanthaster population regulation by a shrimp and a worm. Proceedings from the 4th International Coral Reef Symposium 2, 607–612. Glynn, P.W. 1983. Increased survivorship in corals harboring crustacean symbionts. Marine Biology Letters 4, 105–111. Glynn, P.W. 1987. Some ecological consequences of coral crustacean guard mutualisms in the Indian and Paciic ocean. Symbiosis 4, 301–324. Glynn, P.W. 1988. El Niño warming, coral mortality and reef framework destruction by echinoid bioerosion in the eastern Paciic. Galaxea 7, 129–160. Glynn, P.W. 1994. State of coral reefs in the Galapagos islands: natural versus anthroprogenic impacts. Marine Pollution Bulletin 29, 131–140. Glynn, P.W. 2004. High complexity food webs in low-diversity eastern Paciic Reef coral communities. Ecology 7, 358–367. Glynn, P.W. & D’Croz, l. 1990. Experimental evidence for high temperature stress as the cause of El Niñocoincident coral mortality. Coral Reefs 8, 181–191. Glynn, P.W. & Krupp, D.A. 1986. Feeding biology of a Hawaiian sea star corallivore, Culcita novaeguineae. Journal of Experimental Marine Biology and Ecology 96, 75–96. Glynn, P.W., Perez, M. & Gilchrist, S. 1985. lipid decline in stressed corals and their crustacean symbionts. Biological Bulletin (Woods Hole) 168, 276–284. Glynn, P.W., Stewart, R.H. & McCosker, J.E. 1972. Paciic coral reefs of Panamá: structure, distribution, and predators. Geologische Rundschau 61, 483–519. Glynn P.W., Wellington, G.M. & Wells J.W. 1983. Corals and Coral Reefs in the Galapagos Islands. berkeley, California: university of California Press. Gohar, H.A.F. & Soliman, G.N. 1963. on three mytilid species boring in living corals. Publications of the Marine Biological Station Al-Ghardaga Red Sea 12, 65–98. Goreau, T.F. & Hartman, W.D. 1963. boring sponges as controlling factors in the formation and maintenance of coral reefs. In Mechanisms of Hard Destruction, R.F. Sogrinaes (ed.). Washington, DC: American Association Advancement of Science, 25–54. Goreau, T.F. & Hartman, W.D. 1966. Sponge: effect on the form of reef corals. Science 151, 3708, 343–344. Goreau, T.F., Goreau, N.I., Soot-Ryen, T. & yonge, C.M. 1969. on a new commensal mytilid (Mollusca: bivalvia) opening into the coelenteron of Fungia scufaria (Coelenterata). Journal of Zoology London 158, 171–195. Goreau, T.J. 1992. bleaching and reef community change in Jamaica: 1951–1991. American Zoologist 32, 683–695. 70 CoRAl-ASSoCIATED INvERTEbRATES Goreau, T.J., Hayes, R.l. & McClanahan, T. 2000. Conservation of coral reefs after the 1998 global bleaching event. Conservation Biology 14, 5–15. Gotelli, N.J., Gilchrist, S. & Abele, l.G. 1985. Population biology of Trapezia spp. and other coral-associated decapods. Marine Ecology Progress Series 21, 89–98. Graham, N.A.J. 2007. Ecological versatility and the decline of coral feeding ishes following climate driven coral mortality. Marine Biology 153, 119–127. Graham, N.A.J., Wilson, S.K., Jennings, S., Polunin, N.v.C., bijoux, J.P. & Robinson, J. 2006. Dynamic fragility of oceanic coral reef ecosystems. Proceedings of the National Academy of Sciences of the United States of America 103, 8425–8429. Gratwicke, b. & Speight, M.R. 2005. Effects of habitat complexity on Caribbean marine ish assemblages. Marine Ecology Progress Series 292, 301–310. Gray, J.S. 1997. Marine biodiversity: patterns, threats and conservation needs. Biodiversity and Conservation 6, 153–175. Greenstein, b.J. & Pandoli, J.M. 2008. Escaping the heat: range shift of reef coral taxa in coastal Western Australia. Global Change Biology 14, 513–528. Grifin, S.P., Garcia, R.P. & Weil, E. 2003. bioerosion in coral reef communities in southwest Puerto Rico by the sea urchin Echinometra viridis. Marine Biology 143, 79–84. Hamner, W.M. & Jones, M.S. 1976. Distribution, burrowing, and growth rates of the clam Tridacna crocea on interior reef lats. Oceanologia 24, 207–227. Harmelin-vivien, M.l. & bouchon-Navaro, y. 1983. Feeding diets and signiicance of coral feeding among chaetodontid ishes in Moorea (French Polynesia). Coral Reefs 2, 119–127. Harris, l.G. 1975. Studies on the life history of two coral-eating nudibranchs of the genus Phestilla. Biological Bulletin (Woods Hole) 149, 539–550. Hatcher, b.G. 1988. The primary productivity of coral reefs: a beggar’s banquet. Trends in Ecology and Evolution 3, 106–111. Hayes, J.A. 1990. Distribution, movement and impact of the corallivorous gastropod Coralliophila abbreviata (lamarck) on a Panamanian patch reef. Journal of Experimental Marine Biology and Ecology 142, 25–42. Hernández, l., balart, E.F. & Reyes-bonilla, H. 2009. Checklist of reef decapod crustaceans (Crustacea: Decapoda) in the southern Gulf of California, México. Zootaxa 2119, 39–50. Herring, P.J. 1972. observations on the distribution and feeding habits of some littoral echinoids from Zanzibar. Journal of Natural History 6, 169–175. Hiatt, R.W. & Strasburg, D.W. 1960. Ecological relationships of the ish fauna on coral reefs in the Marshall Islands. Ecological Monographs 30, 65–127. Hixon, M.A. 1997. Effects of reef ishes on corals and algae. In Life and Death of Coral Reefs, C. birkeland (ed.). New york: Chapman and Hall, 230–246. Hixon, M.A. & Menge, b.A. 1991. Species diversity: prey refuges modify the interactive effects of predation and competition. Theoretical Population Biology 39, 178–200. Ho, J., Cheng, y. & Dai, C. 2010. Hastatus faviae n. gen., n. sp., a xariiid copepod parasitic in the honeycomb coral of Taiwan. Crustaceana 83, 89–99. Ho, P.H. & Ng, P.K.l. 2005. on a new species of coral-symbiont crab of the genus Cymo de Haan, 1833 (Crustacea: Decapoda: brachyura: Xanthidae), from the South China Sea. Zootaxa 1029, 31–38. Hobson, E.S. 1974. Feeding relationships of teleostean ishes on coral reefs in Kona, Hawaii. Fishery Bulletin 72, 915–1031. Hoegh-Guldberg, o. 1999. Climate change, coral bleaching and the future of the world’s coral reefs. Marine and Freshwater Research 50, 839–866. Hoeksema, b.W. & best, M.b. 1991. New observations on scleractinian corals from Indonesia: 2. Sipunculanassociated species belonging to the genera Heterocyathus and Heteropsammia. Zoologische Mededelingen 65, 221–245. Holt, R.D. 1987. Prey communications in patchy environments. Oikos 50, 276–290. Holthuis, l.b. 1981. Description of three new species of shrimps (Crustacea, Decopoda, Caridae) from Paciic islands. Proceedings of the Biological Society of Washington 94, 787–800. Houk, P., bograd, S. & van Woesik, R. 2007. The transition zone chlorophyll front can trigger Acanthaster planci outbreaks in the Paciic ocean: historical conirmation. Journal of Oceanography 63, 149–154. 71 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Hsieh, H.J., Chen, C.A., Dai, C., ou, W., Tsai, W. & Su, W. 2007. From the drawing board to the ield: an example for establishing an MPA in Penghu, Taiwan. Aquatic Conservation Marine and Freshwater Ecosystems 17, 619–635. Hughes, T.P. 1994. Catastrophes, phase shifts, and large-scale degradation of a Caribbean coral reef. Science 265, 1547–1551. Hughes, T.P., baird, A.H., bellwood, D.R., Card, M., Connolly, S.R., Folke, C., Grosberg, R., Hoegh-Guldberg, o., Jackson, J.b.C., Kleypas, J., lough, J.M., Marshall, P., Nystrom, M., Palumbi, S.R., Pandoli, J.M., Rosen, b. & Roughgarden, J. 2003. Climate change, human impacts and the resilience of coral reefs. Science 301, 929–933. Hughes, T.P., bellwood, D.R. & Connolly, S.R. 2002. biodiversity hotspots, centres of endemicity, and the conservation of coral reefs. Ecology Letters 5, 775–784. Hughes, T.P., Rodrigues, M.J., bellwood, D.R., Ceccarelli, D., Hoegh-Guldberg, o., McCook, l., Moltschaniwskyj, N., Pratchett, M.S., Steneck, R.S. & Willis, b. 2007. Phase shifts, herbivory, and the resilience of coral reefs to climate change. Current Biology 17, 360–365. Humes, A.G. 1960. New copepods from madreporarian corals. Kieler Meeresforschungen 16, 229–235. Humes, A.G. 1962a. Eight new species of Xariia (Copepoda, Cyclopoida), parasites of corals in Madagascar. Bulletin of the Museum of Comparative Zoology Harvard University 128, 37–63. Humes, A.G. 1962b. Kombia angulata n. gen., n. sp. (Copepoda Cyclopoida) parasitic in a coral in Madagascar. Crustaceana 4, 47–56. Humes, A.G. 1973. Cyclopoid copepods (lichomolgidae) from fungiid corals in New Caledonia. Zoologischer Anzeiger 190, 312–333. Humes, A.G. 1974a. Cyclopoid copepods associated with the coral genera Favia, Favites, Platygyra, and Merulina in New Caledonia. Paciic Science 28, 383–399. Humes, A.G. 1974b. Odontomolgus mundulus n. sp. (Copepoda, Cyclopoida) associated with the scleractinian coral genus Alveopora in New Caledonia. Transactions of the American Microscopical Society 93, 153–162. Humes, A.G. 1978a. lichomolgid copepods (Cyclopoida) associated with fungiid corals (scleractinia) in the Moluccas. Smithsonian Contributions to Zoology 253, 1–48. Humes, A.G. 1978b. lichomolgid copepods (Cyclopoida) associated with the coral genus Montipora in the Moluccas. Publications of the Seto Marine Biological Laboratory 24, 387–407. Humes, A.G. 1978c. A poecilostome copepod parasitic in a scleractinian coral in the Moluccas. Hydrobiologia 58, 119–128. Humes, A.G. 1979a. Coral-inhabiting copepods from the Moluccas, with a synopsis of cyclopoids associated with scleractinian corals. Cahiers de Biologic Marine 20, 77–107. Humes, A.G. 1979b. Poecilostome copepods (lichomolgidae) associated with the scleractinian coral Galaxea in the Moluccas. Journal of Natural History London 13, 507–528. Humes, A.G. 1979c. Cyclopoid copepods (lichomolgidae) associated with the coral genus Favites in the Moluccas. Zoological Journal of the Linnean Society London 66, 95–112. Humes, A.G. 1979d. Cyclopoid copepods (lichomolgidae) associated with the scleractinian Cyphastrea in New Caledonia. Paciic Science 33, 195–206. Humes, A.G. 1981. Harpacticoid copepods associated with Cnidaria in the Indo-West Paciic. Journal of Crustacean Biology 1, 227–240. Humes, A.G. 1984a. Hemicyclops columnaris sp.n. (Copepoda, Poecilostomatoida, Clausidiidae) Associated with a Coral in Panama (Paciic Side). Zoologica Scripta 13, 33–39. Humes, A.G. 1984b. Copepods associated with the scleractinian coral Porites in French Polynesia. Cahiers de Biologie Marine 25, 181–195. Humes, A.G. 1985a. Cnidarians and copepods: a success story. Transactions of the American Microscopical Society 104, 313–320. Humes, A.G. 1985b. Poecilostomatoid copepods parasitic in the scleractinian coral genus Goniastrea in the Moluccas. Publications of the Seto Marine Biological Laboratory 30, 277–286. Humes, A.G. 1985c. A review of the Xariiidae (Copepoda, Poecilostomatoida), parasites of scleractinian corals in the Indo-Paciic. Bulletin of Marine Science 36, 467–632. Humes, A.G. 1986. Two new species of Cerioxynus (Copepoda: Poecilostomatoida) parasitic in corals (Scleractinia: Faviidae) in the South Paciic. Systematic Parasitology 8, 187–198. Humes, A.G. 1991a. Copepoda associated with scleractinian corals on the Great barrier Reef, northeastern Australia, with a key to the genera of the lichomolgidae. Journal of Natural History 25, 1171–1231. 72 CoRAl-ASSoCIATED INvERTEbRATES Humes, A.G. 1991b. Copepoda associated with the scleractinian coral genus Montipora in the Indo-Paciic. Proceedings of the Biological Society of Washington 104, 101–137. Humes, A.G. 1991c. Mandobius regalis gen. et sp. n. (Copepoda: Poecilostomatoida: lichomolgidae) associated with the coral Pectinia lactuca in New Caledonia. Zoologica Scripta 20, 277–282. Humes, A.G. 1992. Copepods (Poecilostomatoida: lichomolgidae) associated with the scleractinian coral Gardineroseris planulata in the Moluccas. Invertebrate Taxonomy 6, 303–335. Humes, A.G. 1993. Poecilostomatoid copepods associated with the scleractinian coral Acropora in the tropical western Paciic ocean. Invertebrate Taxonomy 7, 805–857. Humes, A.G. 1994. How many copepods? Hydrobiologia 292/293, 1–7. Humes, A.G. 1995a. New species of Anchimolgus (Copepoda: Poecilostomatoida: lichomolgidae) associated with the scleractinian coral Goniopora in the southwest Paciic. Journal of Natural History 29, 65–84. Humes, A.G. 1995b. Poecilostomatoid copepods from the coral Leptoria tenuis in New Caledonia. Cahiers de Biologie Marine 36, 69–80. Humes, A.G. 1996a. Anchimolgus gratus n. sp. (Copepoda: Anchimolgidae), associated with the scleractinian coral Lithactinia novaehiberniae in New Caledonia. Smithsonian Contributions to Zoology 66, 193–200. Humes, A.G. 1996b. New genera of Copepoda (Poecilostomatoida) from the scleractinian coral Psammocora in New Caledonia. Zoological Journal of the Linnean Society London 118, 59–82. Humes, A.G. 1997a. Copepoda (Siphonostomatoida) associated with the fungiid coral Parahalomitra in the southwestern Paciic. Journal of Natural History 31, 57–68. Humes, A.G. 1997b. Two new copepod genera (Poecilostomatoida) associated with the scleractinian coral Psammocora in New Caldonia. Zoologica Scripta 26, 51–60. Humes, A.G. & Dojiri, M. 1982. Xariiidae (Copepoda) parasitic in Indo-Paciic scleractinian corals. Beaufortia 32, 139–228. Humes, A.G. & Dojiri, M. 1983. Copepoda (Xariiidae) parasitic in scleractinian corals from the Indo-Paciic. Journal of Natural History 17, 257–307. Humes, A.G. & Frost, b.W. 1964. New lichomolgid copepods (Cyclopoida) associated with alcyonarians and madreporarians in Madagascar. Cahiers de l’Ofice de la Recherche Scientiique et Technique Outre-Mer (ORSTOM), Série Océanographie 6 (Série Nosy-Bé I.I), 131–212. Humes, A.G. & Ho, J.S. 1967. New cyclopoid copepods associated with the coral Psammocora contigua (Esper) in Madagascar. Proceedings of the United States National Museum 122, 1–32. Humes, A.G. & Ho, J.S. 1968. Xariiid copepods (Cyclopoida) parasitic in corals in Madagascar. Bulletin of the Museum of Comparative Zoology Harvard University 136, 415–459. Humes, A.G. & Stock, J.H. 1972. Preliminary notes on a revision of the lichomolgidae, cyclopoid copepods mainly associated with marine invertebrates. Bulletin of the Zoological Museum of the University of Amsterdam 2, 121–133. Humes, A.G. & Stock, J.H. 1973. A revision of the family lichomolgidae Kossman, 1877, cyclopoid copepods mainly associated with marine invertebrates. Smithsonian Contributions to Zoology 127, 1–368. Huston, M. 1985. variation in coral growth rates with depth at Discovery bay, Jamaica. Coral Reefs 4, 19–25. Hutchings, P.A. 1986. biological destruction of coral reefs—a review. Coral Reefs 4, 239–252. Hutchings, P.A. 2008. Role of polychaetes in bioerosion of coral substrate. In Erlangen Earth Conference Series, l. Tapanila & M. Wisshak (eds). New york: Springer, 249–264. Hutchings, P.A. 2011. bioerosion. In Encyclopedia of Modern Coral Reefs-Structure, Form and Processes, D. Hopley (ed.). berlin: Springer-verlag, 139–156. Hutchings, P., Ahyong, S., byrne, M., Przeslawski, R. & Worheide, G. 2007. vulnerability of benthic invertebrates of the Great barrier Reef to climate change. In Climate Change and the Great Barrier Reef: A Vulnerability Assessment, J.E. Johnson & P.A. Marshall (eds). Townsville, Australia: Great barrier Reef Marine Park Authority and Australian Greenhouse ofice, 309–356. Hutchings, P.A. & bamber, l. 1985. variability of bioerosion rates at lizard Island, GbR: preliminary attempts to explain these rates and their signiicance. Proceedings of the Fifth International Coral Reef Congress, Tahiti 5, 333–338. Jackson, J.b.C., Kirby, M.X., berger, W.H., botsford, l.W., bourque, b.J., bradbury, R.H., Cooke, R., Erlandson, J., Estes, J.A., Hughes, T.P., Kidwell, S., lange, C.b., leniham, H.S., Pandoli, J.M., Peterson, C.H., Steneck, R.S., Tegner, M.J. & Warner, R.R. 2001. Historical overishing and the recent collapse of coastal ecosystems. Science 293, 629–637. 73 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Jennings, S., boulle, D.P. & Polunin, N.v.C. 1996. Habitat correlates of the distribution and biomass of Seychelles’ reef ishes. Environmental Biology of Fishes 46, 15–25. Jokiel, P.l. & Townsley, S.J. 1974. biology of the polyclad Prosthiostomum (Prosthiostomum) sp., a new coral parasite from Hawaii. Paciic Science 28, 361–373. Jones, C.G., lawton, J.H. & Shachak, M. 1994. organisms as ecosystem engineers. Oikos 69, 373–386. Jones, D. & Morgan, G. 2002. A Field Guide to Crustaceans of Australian Waters. Sydney: Reed New Holland. Jones, G.P., McCormick, M.I., Srinivasan, M. & Eagle, J.v. 2004. Coral decline threatens ish biodiversity in marine reserves. Proceedings of the National Academy of Sciences of the United States of America 101, 8251–8253. Jones, G.P. & Syms, C. 1998. Disturbance, habitat structure and the ecology of ishes on coral reefs. Australian Journal of Ecology 23, 287–297. Julliard, R., Jiguet, F. & Couvet, D. 2003. Common birds facing global changes: what makes a species at risk? Global Change Biology 10, 148–154. Karlson, R.H. & Hurd, l.E. 1993. Disturbance, coral reef communities, and changing ecological paradigms. Coral Reefs 12, 117–125. Kassen, R. 2002. The experimental evolution of specialists, generalists, and the maintenance of diversity. Journal of Evolutionary Biology 15, 173–190. Keesing, J.K. 1990. Feeding biology of the crown-of-thorns starish, Acanthaster planci (Linnaeus). PhD thesis, James Cook university of North Queensland, Townsville, Australia. Keesing, J.K. 1992. Inluence of persistent sub-infestation density Acanthaster planci (l.) and high density Echinometra mathaei (de blainville) populations on coral reef community structure in okinawa, Japan. Proceedings of the 7th International Coral Reef Symposium 2, 769–779. Kleypas, J.A., Feely, R.A., Fabry, v.J., langdon, C., Sabine, C.l. & Robbins, l.l. 2006. Impacts of ocean acidiication on coral reefs and other marine calciiers: a guide for future research, report of a workshop held 18–20 April 2005, St. Petersburg, Florida, sponsored by NSF, NoAA, and the u.S. Geological Survey. Kim, I.H. 2003. Copepods (Crustacea) associated with marine invertebrates from New Caledonia. Korean Journal of Systematic Zoology Special Issue 4, 1–167. Kim, I.H. 2004. Copepods (Crustacea) associated with marine invertebrates from Great barrier Reef, Australia. Korean Journal of Systematic Zoology 20, 109–140. Kim, I.H. 2005. Four new species of the genus Panjakus (Copepoda, Cyclopoida, Anchimolgidae) associated with scleractinian corals (Cnidaria) from the Moluccas. Integrative Biosciences 9, 215–228. Kim, I.H. 2006. Copepoda (Poecilostomatoida: Anchimolgidae) associated with the scleractinian coral Gardineroseris planulata (Dana) from the Moluccas. Korean Journal of Systematic Zoology 22, 63–78. Kim, I.H. 2007. Copepods (Crustacea) associated with marine invertebrates from the Moluccas. Korean Journal of Systematic Zoology Special Issue 6, 1–126. Kitahara, M., Sei, K. & Fujii, K. 2000. Patterns in the structure of grassland butterly communities along a gradient of human disturbance: further analysis based on the generalist/specialist concept. Population Ecology 42, 135–144. Knudsen, J.W. 1967. Trapezia and Tetralia (Decapoda, brachyura, Xanthidae) as obligate ectoparasites of the pocilloporid and acroporid corals. Paciic Science 21, 50–57. Kohn, A.J. & leviten, P.J. 1976. Effect of habitat complexity on population density and species richness in tropical intertidal predatory gastropod assemblages. Oecologia 25, 199–210. Kokita, T. & Nakazono, A. 2001. Rapid response of an obligately corallivorous ileish Oxymonacanthus longirostris (Monacanthidae) to a mass coral bleaching event. Coral Reefs 20, 155–158. Kropp, R.K. 1984. Tanaocheles stenochilus, a new genus and species of crab from Guam, Mariana Islands (brachyura: Xanthidae). Proceedings of the Biological Society of Washington 97, 744–747. Kropp, R.K. 1989. A revision of the Paciic species of gall crabs, genus Opecarcinus (Crustacea: Cryptochiridae). Bulletin of Marine Science. 45, 98–129. Kropp, R.K. 1990. Revision of the genera of gall crabs (Crustacea: Cryptochiridae) occurring in the Paciic ocean. Paciic Science 44, 417–448. Kropp, R.K. 1994. The gall crabs (Crustacea: Decapoda: brachyura: Cryptochiridae) of the Rumphius expeditions revisited, with descriptions of three new species. Rafles Bulletin of Zoology 42, 521–538. Kropp, R.K. 1995. Lithoscaptus pardalotus, a new species of coral-dwelling gall crab (Crustacea: brachyura: Cryptochiridae) from belau. Proceedings of the Biological Society of Washington 108, 637–642. 74 CoRAl-ASSoCIATED INvERTEbRATES Kropp, R.K. & Manning, R.b. 1987. The Atlantic gall crabs, family Cryptochiridae (Crustacea: Decapoda: brachyura). Smithsonian Contributions to Zoology 462, 1–21. Kropp, R.K. & Manning, R.b. 1996. Crustacea Decapoda: Two new genera and species of deep water gall crabs from the Indo-west Paciic (Cryptochiridae). Mémoires du Muséum National d’Histoire Naturelle 168, 531–539. Kurihara, H., Asai, T., Kato, S. & Ishimatsu, A. 2008. Effects of elevated pCo2 on early development in the mussel Mytilus galloprovincialis. Aquatic Biology 4, 225–233. Kurihara, H. & Shirayama, y. 2004. Effects of increased atmospheric Co2 on sea urchin early development. Marine Ecology Progress Series 274, 161–169. lawson, G.l., Kramer, D.l. & Hunte, W. 1999. Size-related habitat use and schooling behaviour in two species of surgeonish (Acanthurus bahianus and A. coeruleus) on a fringing reef in barbados, West Indies. Environmental Biology of Fishes 54, 19–33. lawton, J.H. 1993. Range, population abundance and conservation. Trends in Ecology and Evolution 8, 409–413. lee, A. & Sin, T.S. 2009. Trapezia septata Dana, 1852 (brachyura, Trapeziidae): a new record for Singapore with notes on its relationship with the host coral, Pocillopora verrucosa. Crustaceana 82, 1603–1608. levin, l.A. 2006. Recent progress in understanding larval dispersal: new directions and digressions. Integrative and Comparative Biology 46, 282–297. lieske, E. & Myers, R. 1994. Coral Reef Fishes—Indo-Paciic and Caribbean. london: Harper Collins. lindahl, u., Öhman, M.C. & Schelten, C.K. 2001. The 1997/1998 mass mortality of corals: effects on ish communities on a Tanzanian coral reef. Marine Pollution Bulletin 2, 127–131. lobban, C. 2002. Ciliate-Symbiodinium symbiosis spotted on reefs. Coral Reefs 21, 332. lough, J.M. & barnes, D.J. 1990. Possible relationships between environmental variables and skeletal density in a coral colony from the central Great barrier Reef. Journal of Experimental Marine Biology and Ecology 134, 221–241. loya, y., Sakai, K., yamazato, K., Nakano, y., Sambali, H. & van Woesik, R. 2001. Coral bleaching: the winners and the losers. Ecology Letters 4, 122–131. luckhurst, b.E. & luckhurst, K. 1978. Analysis of the inluence of substrate variables on coral reef ish communities. Marine Biology 49, 317–323. Macintyre, I.G., Goodbody, I. Riitzler, K., litter, D.S. & litter, M.M. 2000. A general biological and geological survey of the rims of ponds in the major mangrove islands of the Pelican Cays, belize. Atoll Research Bulletin 467, 13–34. Manning, R.b. 1991. Crustacea Decapoda: Cecidocarcinus zibrowii, a new deep-water gall crab (Cryptochiridae) from New Caledonia. Memoirs du Museum National d’Histoire Naturelle 152, 515–520. Marin, I.N. 2008. Description of two new species from the genera Palaemonella Dana, 1852 and Vir Holthuis, 1952 (Crustacea: Caridea: Palaemonidae: Pontoniinae). Zoologische Mededelingen 82, 375–390. Marsden, J.R. 1962. A coral-eating polychaete. Nature 193, 598. Marshall, P.A. & baird, A.H. 2000. bleaching of corals on the Great barrier Reef: differential susceptibilities among taxa. Coral Reefs 19, 155–163. McClanahan, T.R. 1994. Kenyan coral reef lagoon ish: associations with reef managements, substrate complexity, and sea urchins. Coral Reefs 13, 231–241. McClanahan, T.R. 1997. Dynamics of Drupella cornus populations on Kenyan Coral Reefs. Proceedings of the 8th International Coral Reef Symposium 1, 633–638. McClanahan, T.R., baird, A.H., Marshall, P.A. & Toscano, M.A. 2004. Comparing bleaching and mortality responses of hard coral between southern Kenya and the Great barrier Reef, Australia. Marine Pollution Bulletin 48, 327–335. McConnell, o., longley, R.E. & Koehn, F.E. 1994. The discovery of marine natural products with therapeutic potential. In The Discovery of Natural Products with Therapeutic Potential, v.P. Gullo (ed.). boston: butterworth-Heinemann, 109–174. McKinney, M.l. 1997. Extinction vulnerability and selectivity: combining ecological and paleontological views. Annual Review of Ecology and Systematics 28, 495–516. Mclaughlin, P.A. & lemaitre, R. 1993. A review of the hermit crab genus Paguritta (Decapoda: Anomura: Paguridae) with descriptions of three new species. Rafles Bulletin of Zoology 41, 1–29. Menge, b.A. 1976. organization of New England rocky intertidal community—role of predation, competition, and environmental heterogeneity. Ecological Monographs 46, 355–393. 75 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Meyer, J.l. & Schultz, E.T. 1985. Tissue condition and growth rate of corals associated with schooling ish. Limnology and Oceanography 30, 157–166. Mitsuhashi, M. & Takeda, M. 2008. Identity of the coral-associated pontoniine shrimp species, Coralliocaris nudirostris (Heller, 1861) and C. venusta Kemp, 1922 (Crustacea: Decapoda: Palaemonidae), with descriptions of two new species. Zootaxa 1703, 1–24. Mohammed, T.A. & yassien, M.H. 2008. bivalve assemblages on living coral species in the Northern Red Sea, Egypt. Journal of Shellish Research 27, 1271–1223. Mokady, o., Arazi, G., bonar, D.b. & loya, y. 1991. Coral host speciicity in settlement and metamorphosis of the date mussel LIthophaga lessepsiana (vaillant, 1865). Journal of Experimental Marine Biology and Ecology 146, 205–216. Mokady, o., loya, y. & lazar, b. 1998. Ammonium contribution from boring bivalves to their coral host—a mutualistic symbiosis? Marine Ecology Progress Series 169, 295–301. Moore, R.J. 1978. Is Acanthaster planci an r-strategist? Nature 271, 66–57. Moran, P.J. 1986. The Acanthaster phenomenon. Oceanography and Marine Biology An Annual Review 24, 379–480. Morgan, G.J. 1987. Abbreviated development in Paguristes frontalis (Milne Edwards, 1836) (Anomura: Diogenidae) from southern Australia. Journal of Crustacean Biology 7, 536–540. Morgan, G.J. 1990. A collection of Thalassinidea, Anomura and brachyura (Crustacea: Decapoda) from the Kimberley region of north-western Australia. Zoologische Verhandelingen 265, 1–90. Morgan, G.J. 1991. A review of the hermit crab genus Calcinus Dana (Crustacea: Decapoda: Diogenidae) from Australia, with descriptions of two new species. Invertebrate Taxonomy 5, 869–913. Morgan, G.J. 1993. Three new species of Pagurixus (Crustacea, Decapoda, Paguridae) from western Australia, with notes on other Australian species. In Proceedings of the Fifth International Marine Biological Workshop: The Marine Flora and Fauna of Rottnest Island, F.E. Wells et al. (eds). Perth, Australia: Western Australian Museum, 163–181. Morgan, G.J. & Forest, J. 1991. A new genus and species of hermit crab (Crustacea, Anomura, Diogenidae) from the Timor Sea, north Australia. Bulletin du Muséum National d’Histoire Naturelle 13, 189–202. Morton, b., blackmore, G. & Kwok, C.T. 2002. Corallivory and prey choice by Drupella rugosa (Gastropoda: Muricidae) in Hong Kong. Journal of Molluscan Studies 68, 217–223. Morton, b. & blackmore, G. 2009. Seasonal variations in the density of and corallivory by Drupella rugosa and Cronia margariticola (Caenogastropoda: Muricidae) from the coastal waters of Hong Kong: ‘plagues’ or ‘aggregations’? Journal of the Marine Biological Association of the United Kingdom 89, 147–159. Mueter, F.J. & litzow, M.A. 2008. Sea ice retreat alters the biogeography of the bering Sea continental shelf. Ecological Applications 18, 309–320. Muller-Parker, G.I. & Simon, K.D. 2001. Temperate and tropical algal-sea anemone symbioses. Invertebrate Biology 120, 104–123. Munday, P.l. 2004. Habitat loss, resource specialisation, and extinction on coral reefs. Global Change Biology 10, 1642–1647. Munday, P.l., Jones, G.P. & Caley, M.J. 1997. Habitat specialisation and the distribution and abundance of coral-dwelling gobies. Marine Ecology Progress Series 152, 227–239. Munday, P.l., Jones, G.P. & Caley, M.J. 2001. Interspeciic competition and coexistence in a guild of coral dwelling gobies. Ecology 82, 2177–2189. Munday, P.l., Jones, G.P., Sheaves, M., Williams, A.J. & Goby, G. 2007. vulnerability of ishes on the Great barrier Reef to climate change. In Climate Change and the Great Barrier Reef, J. Johnson & P. Marshall (eds). Townsville, Australia: Great barrier Reef Marine Park Authority, 357–392. Naeem, S. & li, S.b. 1997. biodiversity enhances ecosystem reliability. Nature 390, 507–509. Nair, b.u. 1983. on ive new species of xariiid copepods from the Arabian Sea. In Selected Papers on Crustacea, P.A. John (ed.). Trivandrum, India: Prof. N. Krishna Pillai Farewell Committee, 11–25. Nair, b.u. & Pillai, N.K. 1986. Three new species of copepods associated with South Indian invertebrates. Crustaceana Leiden 50, 27–38. Nielsen Tackett, D. & Tackett, l. 2002. Reef Life: Natural History and Behaviors of Marine Fishes and Invertebrates. Neptune City, New Jersey: T.F.H Publications. 76 CoRAl-ASSoCIATED INvERTEbRATES Ng, P.K.l. & Clark, P.F. 2000. The Indo-Paciic Pilumnidae XII. on the familial placement of Chlorodiella bidentata (Nobili, 1901) and Tanaocheles stenochilus Kropp, 1984 using adult and larval characters with the establishment of a new subfamily, Tanaochelinae (Crustacea: Decapoda: brachyura). Journal of Natural History 34, 207–245. Nugues, M.M. & bak, R.P.M. 2009. brown-band syndrome on feeding scars of the crown-of-thorn starish Acanthaster planci. Coral Reefs 28, 507–510. Nyström, M., Folke, C. & Moberg, F. 2000. Coral reef disturbance and resilience in a human-dominated environment. Trends in Ecology and Evolution 15, 413–417. o’Connor, M.I., bruno, J.F., Gaines, S.D., Halpern, b.S., lester, S.E., Kinlan, b.P. & Weiss, J.M. 2007. Temperature control of larval dispersal and the implications for marine ecology, evolution, and conservation. Proceedings of the National Academy of Sciences of the United States of America 104, 1266–1271. ogawa, K., Pillay, R.M. & Kawasaki, H. 1998. Coral-inhabiting barnacles (Cirripedia; Pyrgomatidae) from Albion, west coast of Republic of Mauritius. Bulletin of the Biogeographical Society of Japan 53, 1–21. ogden, J.C. 1977. Carbonate-sediment production by parrot ish and sea urchins on Caribbean reefs. American Association of Petroleum Geologists Studies in Geology 4, 281–288. ogunlana, M.v., Hooge, M.D., benayahu ,y., yonas, y.I., barneah, o. & Tyler, S. 2005. Waminoa brickneri n. sp. (Acoela: Acoelomorpha) associated with corals in the Red Sea. Zootaxa 1008, 1–11. Öhman, M.C. & Rajasuriya, A. 1998. Relationships between habitat structure and ish communities on coral and sandstone reefs. Environmental Biology of Fishes 53, 19–31. okuda, S. 1937. Spioniform polychaetesfrom Japan. Journal of the Faculty of Science Hokkaido Imperial University Series 6 Zoology 5, 217–254. okuno, J. 2004. Periclimenes speciosus, a new species of anthozoan associated shrimp (Crustacea: Decapoda: Palaemonidae) from southern Japan. Zoological Science 21, 865– 875. olivera, b.M., Rivier, J., Clark, C., Ramilo, C.A., Corpuz, G.P., Abogadie, F.C., Mena, E.E., Woodward, S.R., Hillyard, D.R. & Cruz, l.J. 1990. Diversity of Conus neuropeptides. Science 249, 257–63. oliverio, M. 2009. Diversity of Coralliophilinae (Mollusca, Neogastropoda, Muricidae) at Austral Islands (South Paciic). Zoosystema 31, 759–789. oren, u., brickner, I. & loya, y. 1998. Prudent sessile feeding by the corallivore Coralliophila violacea on coral energy sinks. Proceedings of the Royal Society London Series B 265, 2043–2050. orians, G.H. & Wittenberger, J.F. 1991. Spatial and temporal scales in habitat selection. American Naturalist 137, S29–S49. ormond, R.F.G., Hanscomb, N.J. & beach D.H. 1976. Food selection and learning in the crown-of-thorns starish Acanthaster planci (l.). Marine Behavior and Physiology 4, 93–105. otter, G.W. 1937. Rock-destroying organisms in relation to coral reefs. Scientiic Reports/British Museum (National History), Great Barrier Reef Expedition 1928–1929 1, 323–352. Page, C.A. & Willis, b.l. 2007. Epidemiology of skeletal eroding band on the Great barrier Reef and the role of injury in the initiation of this widespread coral disease. Coral Reefs 27, 257–272. Palmer, A.R. 1994. Temperature sensitivity, rate of development, and time to maturity: geographic variation in laboratory reared laboratory reared Nucella and a cross-phyletic overview. In Reproduction and Development of Marine Invertebrates, W.H. Wilson Jr. et al. (eds). baltimore: Johns Hopkins university Press, 177–194. Pandoli, J.M., bradbury, R.H., Sala, E., Hughes, T.P., bjorndal, K.A., Cooke, R.G., McArdle, D., McClenachan, l., Newman, M.J.H., Paredes, G., Warner, R.R. & Jackson, J.b.C. 2003. Global trajectories of the longterm decline of coral reef systems. Science 301, 955–958. Pang, R.K. 1973. The ecology of some Jamaican excavating sponges. Bulletin of Marine Science 23, 227–243. Pari, N., Peyrot–Clausade, M. & Hutchings, P.A. 2002. bioerosion of experimental substrates on high islands and atoll lagoons (French Polynesia) during 5 years of exposure. Journal of Experimental Marine Biology and Ecology 276, 109–127. Patton, W.K. 1966. Decapod Crustacea commensal with Queensland branching corals. Crustaceana 10, 271–295. Patton, W.K. 1974. Community structure among the animals inhabiting the coral Pocillopora damicornis at Heron Island, Australia. In Symbiosis in the Sea, W.b. vernberg (ed.). Columbia, South Carolina: university of South Carolina Press, 219–243. 77 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Patton, W.K. 1976. Animals associated with living corals. In Biology and Geology of Coral Reefs, o.A. Jones & R. Endean (eds). New york: Academic Press, 1–36. Patton, W.K 1994. Distribution and ecology of animals associated with branching corals (Acropora spp.) from the Great barrier Reef; Australia. Bulletin of Marine Science 55, 193–211. Paulay,G. 1997. Diversity and distribution of reef organisms. In Life and Death of Coral Reefs, C. birkeland (ed.). New york: Chapman and Hall, 303–304. Paulay, G., Kropp, R., Ng, P.K.l. & Eldredge, l.G. 2003. The crustaceans and pycnogonids of the Mariana Islands. Micronesica 35–36, 456–513. Pawlik, J.R. 1992. Chemical ecology of the settlement of marine invertebrates. Oceanography and Marine Biology An Annual Review 30, 273–335. Pearson, R.G. & Endean, R. 1969. A preliminary study of the coral predator Acanthaster planci (l.) (Asteroidea) on the Great barrier Reef. Fish Notes 3, 27–55. Pettibone, M.H. 1993. Scaled polychaetes (Polynoidae) associated with ophiuroids and other invertebrates and review of species referred to Malmgrenia McIntosh and replaced by Malmgreniella Hartman, with descriptions of new taxa. Smithsonian Contributions to Zoology 538, 1–9. Peyrot-Clausade, M., Hutchings, P. & Richard, G. 1992. Temporal variations of macroborers in massive Porites lobata on Moorea, French Polynesia. Coral Reefs 11, 161–166. Ponder, W.F., Hutchings, P.A. & Chapman, R. 2002. overview of the conservation of Australia’s marine invertebrates. A Report for Environmental Australia, Australian Museum, Sydney. Available http://www. amonline.net.au/invertebrates/marine_overview/index.html. Accessed June 3, 2010. Porter, J.W. & Targett, N.M. 1988. Allelochemical interactions between sponges and corals. Biological Bulletin (Woods Hole) 175, 230–239. Potts, D.C. 1981. Crown-of-thorns starish man-induced pest or natural phenomenon? In The Ecology of Pests, R.l. Kitching & R.E. Jones (eds). Melbourne: CSIRo Publications Service, 55–86. Poupin, J. & lemaitre, R. 2003. Hermit crabs of the genus Calcinus Dana, 1851 (Decapoda: Anomura: Diogenidae) from the Austral Islands, French Polynesia, with description of a new species. Zootaxa 391, 1–20. Pratchett, M.S. 2001. Inluence of coral symbionts on feeding preferences of crown-of-thorns starish Acanthaster planci in the western Paciic. Marine Ecology Progress Series 214, 111–119. Pratchett, M.S. 2005. Dietary overlap among coral-feeding butterlyishes (Chaetodontidae) at lizard Island, northern Great barrier Reef. Marine Biology 148, 373–382. Pratchett, M.S. 2007. Dietary selection by coral-feeding butterlyishes (Chaetodontidae) on the Great barrier Reef, Australia. Rafles Bulletin of Zoology 14, S161–S166. Pratchett, M.S. 2010. Changes in coral assemblages during an outbreak of Acanthaster planci at lizard Island, northern Great barrier Reef (1995–1999). Coral Reefs 29, 717–725. Pratchett, M.S., Graham, N.A.J., Sheppard, C.R.C. & Mayes, b. 2010. Are infestations of Cymo melanodactylus killing Acropora cytherea in the Chagos archipelago? Coral Reefs doi:10.1007/s00338-010-0654-x. Pratchett, M.S., Munday, P.l., Wilson, S.K., Graham, N.A.J., Cinner, J.E., bellwood, D.R., Jones, G.P., Polunin, N.v.C. & McClanahan, T.R. 2008. Effects of climate-induced coral bleaching on coral-reef ishes—ecological and economical consequences. Oceanography and Marine Biology An Annual Review 46, 251–296. Pratchett, M.S., vytopil, E. & Parkes, P. 2000. Coral crabs inluence the feeding patterns of crown-of-thorns starish. Coral Reefs 19, 118. Pratchett, M.S., Wilson, S.K. & baird, A.H. 2006. Decline in the abundance of Chaetodon butterlyishes following extensive coral depletion. Journal of Fish Biology 69, 1269–1280. Pratchett, M.S., Wilson, S.K., berumen, M.l. & McCormick, M.I. 2004. Sub-lethal effects of coral bleaching on an obligate coral feeding butterlyish. Coral Reefs 23, 352–356. Pratchett, M.S., Wilson, S.K., Graham, N.A.J., Munday, P.l., Jones, G.P. & Polunin, N.v.C. 2009. Coral bleaching and consequences for motile reef organisms: past, present and uncertain future effects. In Coral Bleaching: Patterns, Processes, Causes and Consequences, M.J.H. van oppen & J.M. lough (eds). berlin: Springer-verlag, 139–158. Precht, W.F. & Aronson, R.b. 2004. Climate lickers and range shifts of reef corals. Frontiers in Ecology and the Environment 2, 307–314. 78 CoRAl-ASSoCIATED INvERTEbRATES Preston, N.P. & Doherty, P.J. 1990. Cross-shelf patterns in the community structure of coral-dwelling Crustacea in the central region of the Great barrier Reef. I. Agile shrimps. Marine Ecology Progress Series 66, 47–61. Przeslawski, R., Ahyong, S., byrne, M., Wörheide, G. & Hutchings, P. 2008. beyond corals and ish: the effects of climate change on non-coral benthic invertebrates of tropical reefs. Global Change Biology 14, 1–23. Pulliam, H.R. & Danielson, b.J. 1991. Sources, sinks, and habitat selection: a landscape perspective on population dynamics. American Naturalist 137, S50–S66. Raven, J., Caldeira, K., Elderield, H., Hoegh-Guldberg, o., liss, P., Riebesell, u., Shepherd, J., Turley, C. & Watson, A. 2005. ocean acidiication due to increasing atmospheric carbon dioxide. The Royal Society Policy Document 12/05. Cardiff, Wales: Cloyvedon Press. Ray, G.C. 1985. Man and the sea: the ecological challenge. American Zoologist 25, 451–468. Ray, G.C. & Grassle, J.F. 1991. Marine biological diversity. Bioscience 41, 453–457. Reaka-Kudla, M.l. 1997. The global biodiversity of coral reefs: A comparison with rain forests. In Biodiversity II: Understanding and Protecting our Natural Resources, M.l. Reaka-Kudla et al. (eds). Washington, DC: Joseph Henry/National Academy Press, 83–108. Reese, E.S. 1977. Coevolution of corals and coral feeding ishes of the family Chaetodontidae. In Proceedings of the Third International Coral Reef Symposium, Vol. 1. Miami, Florida: university of Miami, 267–274. Reitzel, A.M., Miner, b.G. & McEdward, l.R. 2004. Relationships between spawning date and larval development time for benthic marine invertebrates: a modelling approach. Marine Ecology Progress Series 280, 13–23. Reyes-bonilla, H. & Calderon-Aguilera, l.E. 1999. Population density, distribution and consumption rates of three corallivores at Cabo Pulmo reef, Gulf of California, Mexico. Marine Ecology 20, 347–357. Ribes, M., Coma, R., Atkinson, M.J. & Kinzie, R.A., III. 2005. Sponges and ascidians control removal of particulate organic nitrogen from coral reef water. Limnology and Oceanography 50, 1480–1489. Richardson, A.J. 2008. In hot water: zooplankton and climate change. ICES Journal of Marine Science 65, 279–295. Richter, C., Wunsch, M., Rasheed, M., Koetter, I. & badran, M.I. 2001. Endoscopic exploration of Red Sea coral reefs reveals dense populations of cavity dwelling sponges. Nature 413, 726–730. Rinkevich, b., Wolodarsky, Z. & loya, y. 1991. Coral-crab association: a compact domain of a multilevel trophic system. Hydrobiologia 216/217, 279–284. Robertson, R. 1970. Review of the predators and parasites of stony corals with special reference to symbiotic prosobranch gastropods. Paciic Science 24, 43–54. Rodríguez-Martínez, R.E., banaszak, A.T. & Jordan-Dahlgren, E. 2001. Necrotic patches affect Acropora palmata (Scleractinia: Acroporidae) in the Mexican Caribbean. Diseases of Aquatic Organisms 47, 229–234. Ross, A. & Newman, W.A. 1969. A coral-eating barnacle. Paciic Science 23, 252–256. Ross, A. & Newman, W.A. 1973. Revision of the coral-inhabiting barnacles (Cirripedia: balanidae). Transactions of the San Diego Society of Natural History 17, 137–174. Ross, A. & Newman, W.A. 2000. Coral barnacles: Cenozoic decline and extinction in the Atlantic/East Paciic versus diversiication in the Indo-West Paciic. Proceedings 9th International Coral Reef Symposium, Bali, Indonesia Vol. 1, 23–27. Ross, A. & Newman, W.A. 2002. A review of the Pyrgoma cancellatum species complex (Cirripedia: Pyrgomatidae). Journal of Natural History 36, 407–421. Rotjan, R.D. & lewis, S.M. 2008. Impact of coral predators on tropical reefs. Marine Ecology Progress Series 367, 73–91. Rudman, W.b. 1981. Further studies on the anatomy and ecology of opisthobranch molluscs feeding on the scleractinian coral Porites. Zoological Journal of the Linnean Society 71, 373–412. Ruppert, E.E., Fox, R.S. & barnes, R.D. 2004. Invertebrate Zoology. A Functional Evolutionary Approach. 7th ed. belmont, California: Thomson brooks/Cole. Rützler, K. 2002. Impact of crustose clionid sponges on Caribbean reef corals. Acta Geologica Hispanica 37, 61–72. Ryer, C.H., van Montfrans, J. & orth, R.J. 1990. utilization of a sea-grass meadow and tidal marsh creek by blue crabs Callinectes sapidus. II. Spatial and temporal patterns of molting. Bulletin of Marine Science 46, 95–104. 79 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Salazar, M.H., Wicksten M.K. & Morrone, J.J. 2005. Redescriptions and taxonomic notes on species of the Synalpheus townsendi Coutière, 1909 complex ((Decapoda: Caridea: Alpheidae). Zootaxa 1027, 1–26. Sale, P.F. 1991. The assembly of reef ish communities: open non-equilibrial systems. In The Ecology of Fishes on Coral Reefs, P.F. Sale (ed.). San Diego, California: Academic Press, 564–598. Sano, M. 1989. Feeding habits of Japanese butterlyishes (Chaetodontidae). Environmental Biology of Fishes 25, 195–203. Sano, M., Shimizu, M. & Nose, y. 1984. Food habits of the teleostean reef ishes in okinawa Isiantd [sic], southern Japan. Bulletin the University Museum University of Tokyo 25, 1–128. Schoener, T.W. 1971. Theory of feeding strategies. Annual Review of Ecology and Systematics 2, 369–404. Schuhmacher, H. 1977. A hermit crab, sessile on corals, exclusively feeds by feathered antennae. Oceanologia 27, 371–3741. Scott, P.J.b. 1980. Associations between scleractinians and coral-boring molluscs in Hong Kong. In Proceedings, First International Workshop on the Malacofauna of Hong Kong and Southern China, b. Morton (ed.). Hong Kong: Hong Kong university Press, 121–138. Scott, P.J.b. 1986. A new species of Lithophaga (bivalvia: lithophaginae) boring corals in the Caribbean. Journal of Molluscan Studies 52, 55–61. Scott, P.J.b. 1987. Associations between corals and macro-infaunal invertebrates in Jamaica, with a list of Caribbean and Atlantic coral associates. Bulletin of Marine Science 40, 271–286. Scott, P.J.b. 1988. Distribution, habitat and morphology of the coral and rock-boring bivalve Lithophaga bisulcata (d’orbigny) (Mytilidae: lithophaginae). Journal of Molluscan Studies 54, 83–85. Sebastian, M.J. & Pillai, N.K. 1973. Humesiella corallicola n.g., n.sp., a cyclopoid copepod associated with coral on the south east coast of India. Hydrobiologia 42, 143–152. Sebens, K.P. 1987. The ecology of indeterminate growth in animals. Annual Review of Ecology and Systematics 18, 371–407. Sebens, K.P. 1994. biodiversity of coral reefs: what are we losing and why? American Zoologist 34, 115–133. Serène, R. 1984. Crustaces Decapodes brachyoures de l’ocean Indien occidental et de la Mer Rouge. Xanthoidea: Xanthidae et Trapeziidae. Addendum Carpiliidae et Menippidae A. Crosnier. Faune Tropicale 24, 1–48. Shannon, T., III & Achatz, J.G. 2007. Convolutriloba macropyga sp. nov., an uncommonly fecund acoel (Acoelomorpha) discovered in tropical aquaria. Zootaxa 1525, 1–17. Shears, N.T. & babcock, R.C. 2002. Marine reserves demonstrate top-down control of community structure on temperate reefs. Oecologia 132, 131–142. Sheppard, C.R.C., Spalding, S., bradshaw, C. & Wilson, S. 2002. Erosion vs. recovery of coral reefs after 1998 El Niño: Chagos reefs, Indian ocean. Ambio 31, 40–48. Sheppard brennand, H., Soars, N., Dworjanyn, S.A., Davis, A.R. & byrne, M. 2010. Impact of ocean warming and ocean acidiication on larval development and calciication in the sea urchin Tripneustes gratilla. PLoS ONE 5, e11372. doi:10.1371/journal.pone.0011372. Shima, J.S., osenberg, C.W. &Stier, A.C. 2010. The vermetid gastropod Dendropoma maximum reduces coral growth and survival. Biology Letters doi:10.1098/rsbl.2010.0291. Shirayama, y. & Horikoshi, M. 1982. A new method of classifying the growth form of corals and its application to a ield survey of coral-associated animals in Kabira Cove, Japan. Journal of the Oceanographical Society of Japan 38,193–207. Shirley, M.A., Hines, A.H. & Wolcott, T.G. 1990. Adaptive signiicance of habitat selection by molting adult blue crabs Callinectes sapidus (Rathbun) within a subestuary of central Chesapeake bay. Journal of Experimental Marine Biology and Ecology 140, 107–119. Simon-blecher, N., Huchon, D. & Achituv, y. 2007. Phylogeny of coral inhabiting barnacles (Cirripedia: Thoracica: Pyrgomatidae) based on 12S, 16S and 18S rDNA analysis. Molecular Phylogenetics and Evolution 44, 1333–1341. Sin, T.M. 1999. Distribution and host specialization in Tetralia crabs (Crustacea: brachyura) symbiotic with corals in the Great barrier Reef, Australia. Bulletin of Marine Science 65, 839–850. Soliman, G.N. 1969. Ecological aspects of some coral-boring gastropods and bivalves of the northwestern Red Sea. American Zoologist 9, 887–894. Spotte, S. 1996. Supply of regenerated nitrogen to sea anemones by their symbiotic shrimp. Journal of Experimental Marine Biology and Ecology 198, 27–36. Stachowicz, J.J. & Hay, M.E. 1999. Mutualism and coral persistence: the role of herbivore resistance to algal chemical defence. Ecology 80, 2085–2101. 80 CoRAl-ASSoCIATED INvERTEbRATES Stella, J.S., Jones, G.P. & Pratchett, M.S. 2010. variation in the structure of epifaunal invertebrate assemblages among coral hosts. Coral Reefs 29, 957–973. Stewart, H.l., Holbrook, S.J., Schmitt, R.J. & brooks, A.J. 2006. Symbiotic crabs maintain coral health by clearing sediments. Coral Reefs 25, 609–615. Stimson, J. 1990. Stimulation of fat-body production in the polyps of the coral Pocillopora damicornis by the presence of mutualistic crabs of the genus Trapezia. Marine Biology 106, 211–218. Stock, J.H. 1975. Corallovexiidae, a new family of transformed copepods endoparasitic in reef corals. Studies of the Fauna of Curaçao and other Caribbean Islands 47, 1–45. Strathmann, R.R., Cameron, R.A. & Strathmann, M.F. 1984. Spirobranchus giganteus (Pallus) breaks a rule for suspension-feeders. Journal of Experimental Marine Biology and Ecology 79, 245–249. Stump, R. 1992. life history characteristics of Acanthaster planci (l.) populations, potential clues to causes of outbreaks. In The Possible Causes and Consequences of Outbreaks of the Crown-of-Thorns Starish, u. Engelhardt & b. lassig (eds). Townsville, Australia: Great barrier Reef Marine Park Authority, 105–116. Suchanek, T.H., Carpenter, R.C., Witman, J.D. & Harvell, C.D. 1983. Sponges as important space competitors in deep Caribbean coral reef communities. In The Ecology of Deep and Shallow Coral Reefs. Symposia Series for Undersea Research 1, M.l. Reaka (ed.). Rockville, Maryland: NoAA/NuRP, 55–60. Sullivan, b., Faulkner, D.J. & Webb, l. 1983. Siphonodictidine, a metabolite of the burrowing sponge Siphonodictyon sp. that inhibits coral growth. Science 221, 1175–1176. Sussman, M., loya, y., Fine, M. & Rosenberg, E. 2003. The marine ireworm Hermodice carunculata is a winter reservoir and spring-summer vector for the coral bleaching pathogen Vibrio shiloi. Environmental Microbiology 5, 250–255. Sweatman, H.P.A. 1995. A ield study of ish predation on juvenile crown-of-thorns starish. Coral Reefs 14, 47–53. Sweatman, H.P.A. 2008. No-take reserves protect coral reefs from predatory starish. Current Biology 18, R598-R599. Takeda, M. & Tamura, y. 1980. Coral-inhabiting crabs of the family Hapalocarcinidae from Japan. v. Genus Cryptochirus. Researches on Crustacea 10, 45–56. Takeda, M. & Tamura, y. 1981. Coral-inhabiting crabs of the family Hapalocarcinidae from Japan. vIII. Genus Pseudocryptochirus and two new genera. Bulletin of the Biogeographical Society of Japan 36, 14–27. Takeda, M. & Tamura, y. 1983. Coral-inhabiting crabs of the family Hapalocarcinidae from Japan. IX. A small collection made at Kushimoto and Koza, the Kii Peninsula. Bulletin of the National Science Museum Tokyo Series A (Zoology) 9, 1–11. Taylor, J.D. 1978. Habitats and diet of predatory gastropods at Addu Atoll, Maldives. Journal of Experimental Marine Biology and Ecology 31, 83–103. ten Hove, H.A. 1994. Serpulidae (Annelida: Polychaeta) from the Seychelles and Amirante Islands. In Oceanic Reefs of the Seychelles. Cruise Reports Netherlands Indian Ocean Program, II, J. van der land (ed.). leiden, The Netherlands: National Museum of Natural History of leiden, 107–116. Tewksbury, J.J., Huey, R.b. & Deutsch, C.A. 2008. Putting the heat on tropical animals. Science 320, 1296–1297. Thomassin, b.A. 1976. Feeding behaviour of the felt-, sponge-, and coral-feeder sea stars, mainly Culcita schmideliana. Helgoländer Wissenschaften Meeresuntersuchungen 28, 51–65. Trautwein, S.E. 2007. Four new species of coral crabs belonging to the genus Tetralia Dana, 1851 (Crustacea, Decapoda, brachyura, Tetraliidae). Zootaxa 1450, 1–20. Trench, R.K. 1993. Microalgal-invertebrate symbioses: a review. Endocytobiosis Cell Research 9, 135–175. Trench, R.K. & Winsor, H. 1987. Symbiosis with dinolagellates in two pelagic latworms, Amphiscolops sp. and Haplodiscus sp. Symbiosis 3, 1–22. Tsuchiya, M., yamauchi, y., Moretzsohn, F. & Tsukiji, M. 1992. Species composition and some population traits of obligate symbiotic xanthid crabs, Trapezia and Tetralia, associated with bleached corals. Proceedings of the 7th International Coral Reef Symposium 1, 56–63. Tsuchiya, M. & yonaha, C. 1992. Community organisation of associates of the scleractinian coral Pocillopora damicornis: effects of colony size and interactions among the obligate symbionts. Galaxea 11, 29–56. Turner, S.T. 1994. Spatial variability in the abundance of the corallivorous gastropod Drupella cornus. Coral Reefs 13, 41–48. 81 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Tyler, J.C. 1971. Habitat preferences of the ishes that dwell in shrub corals on the Great barrier Reef. Proceedings of the National Academy of Sciences of the United States of America 123, 1–26. vannini, M. 1985. A shrimp that speaks crab-ese. Journal of Crustacean Biology 5, 160–167. vazquez, D.P. & Simberloff, D. 2002. Ecological specialization and susceptibility to disturbance: conjectures and refutations. American Naturalist 159, 606–623. veron, J.E.N. 2000. Corals of the World. 3rd ed. Townsville, Australia: Australian Institute of Marine Science. vogler, C., benzie, J., lessios, H., barber, P. & Worheide, G. 2008. The threat to coral reefs multiplied? Four species of crown-of-thorns starish. Biology Letters 4, 696–699. vytopil, E. & Willis, b.l. 2001. Epifaunal community structure in Acropora spp. (Scleractinia) on the Great barrier Reef: Implications of coral morphology and habitat complexity. Coral Reefs 20, 281–288. Walther, G.R., Post, E., Convey, P., Menzel, A., Parmesan, C., beebee, T.J.C., Fromentin, J., Hoegh-Guldberg, o. & barlein, F. 2002. Ecological responses to recent climate change. Nature 416, 389–395. West, J.M. & Salm, R.v. 2003. Resistance and resilience to coral bleaching: implications for coral reef conservation and management. Conservation Biology 17, 956–957. Wilkinson, C.R. 1983. Role of sponges in coral reef structural processes. In Perspectives on Coral Reefs, D.J. barnes (ed.). Townsville, Australia: Australian Institute of Marine Science, 263–274. Wilkinson, C.R. 1999. Global and local threats to coral reef functioning and existence: review and predictions. Marine and Freshwater Research 8, 867–878. Wilkinson, C.R. & buddemeier, R.W. 1994. Global climate change and coral reefs: implications for people and reefs. Report of the uNEPIoC-ASPEI-IuCN Global Task Team on Coral Reefs. Gland, Switzerland: IuCN. Williams, D.E. & Miller, M.W. 2005. Coral disease outbreak: pattern, prevalence and transmission in Acropora cervicornis. Marine Ecology Progress Series 301, 119–128. Williams, D.M. 1986. Temporal variation in the structure of reef slope ish communities (central Great barrier Reef): short term effects of Acanthaster planci infestations. Marine Ecology Progress Series 28, 157–164. Williamson, M. 1997. Marine biodiversity in its global context. In Marine Biodiversity: Patterns and Processes, R.F.G. ormond et al. (eds). Cambridge, uK: Cambridge university Press, 1–17. Wilson, b.R. 1979. A revision of Queensland lithophagine mussels (bivalvia, Mytilidae, lithophaginae). Records of the Australian Museum 32, 435–489. Wilson, S.K., Graham, N.A.J. & Polunin, N.v.C. 2007. Appraisal of visual assessments of habitat complexity and benthic composition on coral reefs. Marine Biology 151, 1069– 1076. Wilson, S.K., Graham, N.A.J., Pratchett, M.S., Jones, G.P. & Polunin, N.v.C. 2006. Multiple disturbances and the global degradation of coral reefs: are reef ishes at risk or resilient? Global Change Biology 12, 2220–2234. Winsor, l. 1990. Marine Turbellaria (Acoela) from North Queensland. Memoirs of the Queensland Museum 28, 785–800. Wood, H.l., Spicer, J.I. & Widdicombe, S. 2008. ocean acidiication may increase calciication rates, but at a cost. Proceedings of the Royal Society B 275, 1767–1773. yamaguchi, M. 1975. Coral-reef asteroids of Guam. Biotropica 7, 12–23. yamaguchi, M. 1986. Acanthaster planci infestations of reefs and coral assemblages in Japan: a retrospective analysis of control efforts. Coral Reefs 5, 23–30. yokochi, H. 2004. Predation damage to corals. In Coral Reefs of Japan. Tokyo: Ministry of the Environment and Coral Reef Society, 49–55. yonge, C.M. 1967. observations on Pedum spondyloideum (Chemnitz) Gmelin, a scallop associated with reefbuilding corals. Proceedings of the Malacological Society of London 37, 311–323. Zann, l., brodie, J.E. & vuki, v. 1990. History and dynamics of the crown-of-thorns starish Acanthaster planci (l.) in the Suva area, Fiji. Coral Reefs 9, 135–144. Zibrowius, H., Southward, E.C. & Day J.H. 1975. New observations on a little-known species of Lumbrineris (Polychaeta) living on various Cnidarians, with notes on its recent and fossil Scleractinian hosts. Journal of the Marine Biological Association of the United Kingdom 55, 83–108. Zuschin, M., Hohenegger, J. & Steininger, F.F. 2001. Molluscan assemblages on coral reefs and associated hard substrata in the northern Red Sea. Coral Reefs 20, 107–116. 82 CoRAl-ASSoCIATED INvERTEbRATES Appendix List of invertebrate species known to utilize coral based on published studies* Higher Taxa Species Coral taxa Type use Reference Gammaropsis sp. Gammaropsis sp. Monp Monp o o H H bergsma 2009 bergsma 2009 Parapilumnus sp. Poc u u black & Prince 1983 Alpheus bicostatus Alpheus bidens Alpheus bucephaloides Alpheus clypeatus Alpheus collumianus Alpheus diadema Alpheus frontalis Alpheus gracilis Alpheus leviusculus Alpheus lottini Alpheus malleodigitus Alpheus obesomanus Alpheus pachychirus Alpheus panamensis Alpheus parvirostris Alpheus strenuus Alpheus sublucanus Alpheus sulcatus Alpheus ventrosus Athanas areteformis Athanas dimorphus Athanas granti Athanas sibogae Pomagnathus corallinus Racilius compressus Synalpheus biungulculatus Synalpheus brevispinus Synalpheus charon Synalpheus digueti Synalpheus mexicanus Synalpheus nobilii Poc Poc Sty Poc Poc Poc, Sty Poc Sty, Poc Poc, Ser, Acr Poc Ser Scl Poc Poc Sty Poc Sty Poc Poc Poc Poc Poc Poc Poc Gal Poc Scl Poc, Sty Poc Poc Poc u F u u u u u u u o u u u u u u u u u u u u u o o u u o u u u u H H H u H H H u F u H u H H u H H H u u u u H H H H H H H H black & Prince 1983 Debelius 2001 Edwards & Emberton 1980 bowers 1970 black & Prince 1983 Austin et al. 1980 Austin et al. 1980 Edwards & Emberton 1980 Stella et al. 2010 Patton 1974 Stella et al. 2010 bruce 1984 black & Prince 1983 Abele & Patton 1976 Edwards & Emberton 1980 black & Prince 1983 Edwards & Emberton 1980 Hernández et al. 2009 Patton 1974 black & Prince 1983 black & Prince 1983 black & Prince 1983 black & Prince 1983 Anker et al. 2006 bruce 1972a Abele & Patton 1976 Salazar et al. 2005 Debelius 2001 Abele & Patton 1976 Abele & Patton 1976 Hernández et al. 2009 Phylum Arthropoda Class: Malacostraca order: Amphipoda Family: Photidae order: Decapoda Family: Acidopsidae Family : Alpheidae continued * Key abbreviations appear on page 104. 83 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Family: Cryptochiridae Species Coral taxa Synalpheus sanjosei Synalpheus sanlucasei Synalpheus tumidomanus Cecidocarcinus brychius Cecidocarcinus zibrowii Cryptochirus coralliodytes Cryptochirus planus Cryptochirus rubrilineatus Dacryomaia edmonsoni Dacryomaia japonica Poc Poc Poc Den Den Fav, ocu Fav, ocu Fav, ocu Tham, Sid, Fav Tham, Sid, Fav u u u o o o o o o o H H u H H H H H H H Hernández et al. 2009 Abele & Patton 1976 black & Prince 1983 Kropp & Manning 1987 Manning 1991 Kropp 1990 Takeda & Tamura 1983 Fize & Serène 1957 Kropp 1990 Takeda & Tamura 1981 Dacryomaia sp. A Dacryomaia sp. b Detocarcinus balssi Sid Sid Rhi, ocu, Cary, Den Mus Mus Mus Mus Mus Fung Fung Poc, Acr, Ser, Sty, Por br. Fav, Mer Fav, Mer Fav, Mer Fav, Mer Fav, Mer Fav, Mer Fav, Mer Fav, Mer Fav, Mer Pav Den, Tub Den, Tub o o o H H H Paulay et al. 2003 Paulay et al. 2003 Kropp & Manning 1987 o o o o o o o o H H H H H H H H Kropp 1990 Takeda & Tamura 1980 Kropp 1994 Fize & Serène 1956 Kropp 1994 Fize & Serène 1956 Kropp 1990 Fize & Serène 1957 o o o o o o o o o o o o H H H H H H H H H H H H Fize & Serène 1956 Takeda & Tamura 1983 Fize & Serène 1957 Fize & Serène 1957 Edmondson 1933 Fize & Serène 1957 Kropp 1995 Kropp 1994 Fize & Serène 1956 Kropp & Manning 1996 Fize & Serène 1956 Fize & Serène 1957 Aga, Pav Aga, Pav Aga, Pav Aga, Sid Aga, Pav Aga, Pav Aga, Pav Aga, Pav Fav Den, Tub Aga, Sid o o o o o o o o o o o H H H H H H H H H H H Kropp 1989 Kropp 1989 Kropp 1989 Fize & Serène 1957 Kropp 1989 Kropp 1989 Kropp 1989 Kropp 1989 Edmondson 1933 Fize & Serène 1957 Fize & Serène 1957 Fizesereneia heimi Fizesereneia ishikawai Fizesereneia latisella Fizesereneia stimpsoni Fizesereneia tholia Fungicola fagei Fungicola utinomi Haplocarcinus marsupialis Hiroia krempi Lithoscaptus grandis Lithoscaptus helleri Lithoscaptus nami Lithoscaptus paciicus Lithoscaptus paradoxus Lithoscaptus pardalotus Lithoscaptus prionotus Lithoscaptus tri Luciades agana Neotroglocarcinus dawydofi Neotroglocarcinus hongkongensis Opecarcinus aurantius Opecarcinus crescentus Opecarcinus granulatus Opecarcinus hypostegus Opecarcinus lobifrons Opecarcinus peliops Opecarcinus pholeter Opecarcinus sierra Pelycomaia minuta Pseudocryptochirus viridis Pseudohapalocarcinus ransoni 84 Type use Reference CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa Family: Diogenidae Family: Domeciidae Family: Dromiidae Family: Epialtidae Family: Galatheidae Family: Gnathophyllidae Family: Hippolytidae Species Coral taxa Sphenomaia pyriforma Troglocarcinus corallicola Fav Ast, Sid, Fav, ocu, Mea, Mus, Cary Poc Fav, Mer, Pec Fav, Mer, Pec Fav, Mer, Pec Phyl Poc Scl Poc Poc Poc Acr, Por br., Poc, Sty, Ser Poc Scl Sty Scl Sty, Acr Poc Scl Sty Scl Scl Scl Poc Acr Acr Acr, Poc Poc, Sty Fav, Pav, Por Scl Sty Poc Poc Poc Sty Sty, Acr, Poc Sty Sty Poc Poc Poc Poc Utinomiella dimorpha Xynomaia boissini Xynomaia sheni Xynomaia verrilli Zibrovia galea Aniculus elegans Calcinus albengai Calcinus californiensis Calcinus explorator Calcinus gouti Calcinus guamensis Calcinus haigae Calcinus inconspicuus Calcinus latens Calcinus lineapropodus Calcinus minutus Calcinus obscurus Calcinus pulcher Calcinus rosaceus Calcinus spicatus Ciliopagurus strigatus Diogenes serenei Trizopagurus magniicus Domecia acanthophora Domecia africana Domecia glabra Domecia hispida Jonesius triunguiculatus Palmyria palmyrensis Cryptodromia granulata Herbstia tumida Menaethius monoceros Pelia paciica Perinea tumida Tylocarcinus styx Galathea afinis Galathea humilis Gnathophyllum panamense Hymenocera picta Hippolysmata vittata Hippolyte varians Type use Reference o o H H Edmondson 1933 Kropp & Manning 1987 o o o o o F u u u F u H H H H H F u u u u u Takeda & Tamura 1980 Fize and Serène 1956 Fize & Serène 1957 Fize & Serène 1957 Kropp & Manning 1996 Glynn et al. 1972 Poupin & lemaitre 2003 Hernández et al. 2009 Hernández et al. 2009 Poupin & lemaitre 2003 Morgan 1991 F u u u u u u u F F u F F F F F F o u u u u u u u u u o F F u u u u u u u u u u u F H H H H H H u u u u u u u u u H u u Poupin & lemaitre 2003 Morgan 1991 Edwards & Emberton 1980 Morgan & Forest 1991 Edwards & Emberton 1980 Abele & Patton 1976 Morgan 1991 Edwards & Emberton 1980 Poupin & lemaitre 2003 Fossa & Nilsen 2000 Morgan 1987 Glynn et al. 1972 Castro et al. 2004 Castro et al. 2004 Castro et al. 2004 Edwards & Emberton 1980 Galil & Takeda 1986 Serène 1984 Edwards & Emberton 1980 Abele & Patton 1976 black & Prince 1983 Abele & Patton 1976 Edwards & Emberton 1980 Edwards & Emberton 1980 Edwards & Emberton 1980 Edwards & Emberton 1980 Abele & Patton 1976 Debelius 2001 Preston & Doherty 1990 Preston & Doherty 1990 continued 85 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Family: Hymenosomatidae Family: leucosiidae Family: Mithracidae Family: Paguridae Family: Palaemonidae Species Coral taxa Hippolyte ventricosus Latreutes mucronatus Lysmata californica Phycocaris simulans Saron marmoratus Thor algicola Thor amboinensis Thor cordelli Thor maldivensis Thor paschalis Poc Poc Poc Poc Poc, Sty Poc Poc Poc Poc Poc F F u F F u F u F F u u u u u u u u u u Preston & Doherty 1990 Preston & Doherty 1990 Abele & Patton 1976 Preston & Doherty 1990 Preston & Doherty 1990 Hernández et al. 2009 Preston & Doherty 1990 Hernández et al. 2009 Preston & Doherty 1990 Preston & Doherty 1990 Elamena abrolhosensis Halicarcinus ovatus Uhlias ellipticus Poc Poc Poc u u u u u u black & Prince 1983 black & Prince 1983 Abele & Patton 1976 Mithraculus forceps Mithrax pygmaeus Teleophrys cristulipes Paguritta sp. Paguritta corallicola Paguritta gracilipes Paguritta harmsi Paguritta kroppi Paguritta morgani Paguritta scottae Pagurixis amsa Pagurus lepidus Brachycarpus biunguiculatus Coralliocaris brevirostris Coralliocaris graminea Coralliocaris labyrintha Coralliocaris macrophthalma Coralliocaris nudirostris Coralliocaris pavoni Coralliocaris sandyi Coralliocaris superba Coralliocaris taiwanensis Coralliocaris venusta Coralliocaris viridis Fennera chacei Harpiliopsis beaupresii Harpiliopsis depressa Harpiliopsis spinigera Harpilius bayeri Harpilius consobrinus Harpilius lutescens Ischnopontonia lophos Jacoste japonica Jacoste lucina ocu Poc Poc Monp Acr Scl Scl Scl Monp Por Poc Poc Poc Acr Acr Acr Acr Acr Pav Acr Acr Pav Acr Acr Poc Poc, Sty Poc, Sty Poc, Sty Poc Poc Acr, Poc Gal Acr Acr F u u F o u u u u u u u u u o u u o u u o o o o o u o u o o o o u u H u u H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H Stachowicz & Hay 1999 Abele & Patton 1976 Abele & Patton 1976 Neilsen Tackett &Tackett 2002 Mclaughlin & lemaitre 1993 Schuhmacher 1977 Mclaughlin & lemaitre 1993 Mclaughlin & lemaitre 1993 Mclaughlin & lemaitre 1993 Mclaughlin & lemaitre 1993 Morgan 1993 Abele & Patton 1976 Hernández et al. 2009 bruce 1977 vytopil & Willis 2001 Mitsuhashi & Takeda 2008 bruce 1977 vytopil & Willis 2001 bruce 1972b Mitsuhashi & Takeda 2008 Patton 1994 Fujino & Miyake 1972 vytopil & Willis 2001 bruce 1974a Gotelli et al. 1985 Edwards & Emberton 1980 Preston & Doherty 1990 Abele & Patton 1976 Holthuis 1981 bruce 1976 Degrave 2000 bruce 1966 Patton 1994 Patton 1994 86 Type use Reference CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa Species Coral taxa Kemponia amymone Acr, Poc, Ser, Sty Fung Goni Poc Poc Eup, Poc, Ser Poc Poc, Ser Acr Sty Poc Poc Sty Poc Por br. Acr, Poc, Por br., Pav Poc Poc Poc Poc Poc Cary Poc, Acr, Ser Sty Poc Sty Scl Poc Poc Poc Acr Acr Poc Gal Tub Den Den Acr Poc, Sty Cary Poc Ple Sty Poc Poc Kemponia kororensis Metapontonia fungiacola Palaemonella assymetrica Palaemonella holmesi Palaemonella rotumana Palaemonella spinulata Palaemonella tenuipes Paratypton siebenrocki Periclimenaeus arabicus Periclimenaeus hectat Periclimenes andamanensis Periclimenes calmani Periclimenes consobrinus Periclimenes dificilus Periclimenes diversipes Family: Pandalidae Periclimenes elegans Periclimenes grandis Periclimenes holthuisi Periclimenes longirostris Periclimenes madreporae Periclimenes magniicus Periclimenes mahei Periclimenes pettihouarsi Periclimenes seychellensis Periclimenes sibogae Periclimenes speciosus Periclimenes spiniferus Periclimenes suvadivensis Periclimenes toloensis Philarius gerlachi Philarius imperialis Philocheras sp. Platycaris latirostris Pontonides maldivensis Pontonides sibogae Pontonides unciger Tectopontonia maziwiae Thaumatocaris streptopus Vir longidactylus Vir orientalis Vir philippinensis Yemenicaris trullicauda Chlorocurtis jactans Chlorotocella gracilis Type use Reference o H Patton 1966 o u u u F F u o u F F u u u u H H H H H H H H H H H H H H H bruce 1977 bruce 1974b Abele & Patton 1976 Hernández et al. 2009 bruce 1972b Preston & Doherty 1990 bruce 1972b bruce 1978 bruce 1974c Austin et al. 1980 Preston & Doherty 1990 Edwards & Emberton 1980 bruce 1974d bruce 1977 bruce 1976 F F F F o u u u F u u u F F u u F u o o o u F o u o u F F H H H H H H H H H H H H H H H H H H H H H H H H H H H u u Preston & Doherty 1990 Preston & Doherty 1990 Preston & Doherty 1990 Preston & Doherty 1990 Preston & Doherty 1990 bruce 1979 bruce 1969 Edwards & Emberton 1980 Preston & Doherty 1990 Edwards & Emberton 1980 okuno 2004 Patton 1974 Preston & Doherty 1990 Preston & Doherty 1990 Patton 1994 Patton 1994 Preston & Doherty 1990 bruce 1978 bruce 2005 bruce 2005 bruce 1978 bruce 1973 Preston & Doherty 1990 Marin 2008 bruce 1972b Neilsen Tackett &Tackett 2002 bruce 1997 Preston & Doherty 1990 Preston & Doherty 1990 continued 87 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Species Coral taxa Family: Pilumnidae Tanaocheles stenochilus Tanaocheles bidentata Pilumnus reticulatus Pilumnus stimpsonii Megalobrachium erosum Megalobrachium smithii Megalobrachium tuberculipes Pachycheles biocellatus Pachycheles granti Pachycheles pisiodes Pachycheles sculptus Pachycheles vicarius Petrolisthes agassizii Petrolisthes carinipes Petrolisthes edwardsii Petrolisthes galanthinus Petrolisthes glasselli Petrolisthes haigae Petrolisthes hirtispinosus Petrolisthes polymitus Petrolisthes sp. Pisidia inaequalis Pisidia magdalenensis Ulloaia perpusillia Thalamitoides tridens Thalamita sp. Processa australiensis Sicyonia sp. Stenopus hispidus Poc Poc Poc Poc Poc Poc Poc u u u u u u u H H H H u u u Kropp 1984 Ng & Clark 2000 Abele & Patton 1976 Abele & Patton 1976 Abele & Patton 1976 Abele & Patton 1976 Abele & Patton 1976 Poc Poc Poc Poc Poc Poc Sty Poc Poc Poc Poc Poc Poc Poc Sty Poc Poc Sty Poc Poc Poc Scl u u u u u u u u u u u u u u u u u u u F F F u u u u u u u u u u u u u u u u u u u u u H Abele & Patton 1976 Austin et al. 1980 Austin et al. 1980 Austin et al. 1980 Abele & Patton 1976 Abele & Patton 1976 Edwards & Emberton 1980 Abele & Patton 1976 Abele & Patton 1976 Hernández et al. 2009 Abele & Patton 1976 Hernández et al. 2009 Abele & Patton 1976 Austin et al. 1980 Edwards & Emberton 1980 Abele & Patton 1976 Abele & Patton 1976 Edwards & Emberton 1980 black & Prince 1983 Preston & Doherty 1990 Preston & Doherty 1990 Jones & Morgan 2002 Neaxius vivesi Poc u u Hernández et al. 2009 Tetralia aurantistellata Tetralia brengelae Tetralia brunalineata Tetralia cavimana Tetralia cinctipes Tetralia glaberrima Tetralia muta Tetralia nigrolineata Tetralia ocucaerulea Tetralia rubridactyla Tetraloides heterodactyla Tetraloides nigrifrons Quadrella boopsis Trapezia areolata Trapezia bella Acr Acr Acr Acr Acr Acr Acr Acr Acr Acr, Poc Acr Acr Den Poc Poc o o o o o o o o o o o o o o o F F F F F F F F F F F F H F F Trautwein 2007 Trautwein 2007 Trautwein 2007 Trautwein 2007 vytopil & Willis 2001 vytopil & Willis 2001 Galil & Clark 1988 vytopil & Willis 2001 Trautwein 2007 Chang et al. 1987 Trautwein 2007 Trautwein 2007 Castro et al. 2004 Austin et al. 1980 Castro et al. 2004 Family: Porcellanidae Family: Portunidae Family: Processidae Family: Sicyoniidae Family: Stenopodidae Family: Strahlaxiidae Family: Tetraliidae Family: Trapeziidae 88 Type use Reference CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa Family: upogebiidae Family: Xanthidae Species Coral taxa Type use Reference Trapezia bidentata Trapezia cheni Trapezia corallina Trapezia cymodoce Trapezia digitalis Trapezia lavopunctata Trapezia ferruginea Trapezia formosa Trapezia garthi Trapezia globosa Poc Poc Poc Poc, Sty Poc, Sty Poc Poc, Sty Poc Poc Poc o o o o o o o o o o F F F F F F F F F F Castro et al. 2004 Chang et al. 1987 Castro et al. 2004 Chang et al. 1987 Chang et al. 1987 Castro et al. 2004 Edwards & Emberton 1980 Chang et al. 1987 Chang et al. 1987 Castro et al. 2004 Trapezia guttata Trapezia intermedia Trapezia lutea Trapezia plana Trapezia neglecta Trapezia punctimanus Trapezia richtersi Trapezia rufopunctata Trapezia septata Trapezia serenei Trapezia speciosa Trapezia tigrina Upogebia operculata Pomatogebia rugosa Chlorodiella laevissima Chlorodiella nigra Chlorodiella spinipes Cyclodius nitidus Cyclodius ungulatus Cycloxanthus bocki Cycloxanthus vittatus Cymo andreossyi Cymo barunae Cymo cerasma Cymo deplanatus Cymo lanatopodus Cymo melanodactylus Cymo quadrilobatus Cymo tuberculatus Etisus anaglyptus Etisus electra Heteractaea lunata Poc, Sty Poc, Sty Poc Poc Poc Poc Poc Poc Poc Poc Poc Poc Por ma. Por ma. Poc, Ser, Acr Poc, Sty Sty Sty Poc Poc Poc Poc Acr Poc Acr Poc Acr Poc Poc Poc Sty Poc o o o o o o o o o o o o o u u u u u u u u u u o u o o o o u u u F F F F F F F F F F F F H H u H H H H H H H H H H H F H H u H H Castro et al. 2004 Castro et al. 2004 Castro et al. 2004 Castro et al. 2004 Castro et al. 2004 Castro et al. 2004 Castro et al. 2004 Chang et al. 1987 Chang et al. 1987 Castro et al. 2004 Castro et al. 2004 Castro et al. 2004 Scott 1987 Fonseca & Cortés 1998 Stella et al. 2010 Austin et al. 1980 Edwards & Emberton 1980 Edwards & Emberton 1980 Patton 1974 Abele & Patton 1976 Abele & Patton 1976 Patton 1974 Ho & Ng 2005 Morgan 1990 Patton 1994 Galil & vannini 1990 Patton 1994 Galil & vannini 1990 Serène 1984 black & Prince 1983 Edwards & Emberton 1980 Abele & Patton 1976 Liocarpilodes integerrimus Liomera rugata Lophoxanthus lamellipes Macromedaeus nudipes Paractaea retusa Sty Sty Poc Ser Sty u u u u u H H H u H Edwards & Emberton 1980 Edwards & Emberton 1980 Abele & Patton 1976 Stella et al. 2010 Edwards & Emberton 1980 continued 89 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa order: Stomatopoda Family: Gonodactylidae Species Coral taxa Type use Reference Paractaea rufopunctata Paraxanthis elegans Psaumis cavipes Pseudoliomera granosimana Pseudoliomera remota Pseudoliomera speciosa Stenorynchus debilis Poc Poc Poc Sty Sty Sty Poc u u u u u u u u H u H H H H black & Prince 1983 black & Prince 1983 black & Prince 1983 Edwards & Emberton 1980 Edwards & Emberton 1980 Edwards & Emberton 1980 Abele & Patton 1976 Gonodactylus falcatus Gonodactylus sp. Sty Poc F u H H Edwards & Emberton 1980 black & Prince 1983 Armatobalanus allium Ahoekia chuangi Ahoekia microtrema Ahoekia tanabensis Arossella lynnae Australhoekia cardenae Cantellius acutum Cantellius albus Cantellius alphonsei Cantellius arcuatum Cantellius brevitergum Cantellius cardenae Cantellius euspinulosa Cantellius gregarius Cantellius hiroi Cantellius hoegi Cantellius iwayama Cantellius iwayama Cantellius madreporae Cantellius octavus Cantellius pallidus Cantellius preobrazhenskyi Cantellius pseudopallidum Cantellius quintus Cantellius secundus Cantellius septimus Cantellius sextus Cantellius sinensis Cantellius sumbawae Cantellius transversalis Cantellius tredecimus Ceratoconcha domingensis Ceratoconcha loridana Ceratoconcha paucicostata Mont Scl Scl Scl Scl Scl Acr Scl Monp Scl Scl Acr Ast Scl Scl Pach Ast Ast Scl Scl Ast Scl Scl Scl Acr Monp Scl Scl Scl Scl Ast Scl Scl Scl o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H Simon-blecher et al. 2007 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Achituv & Hoeksema 2003 Ross & Newman 2000 Achituv 2001 ogawa et al. 1998 ogawa et al. 1998 Achituv & Hoeksema 2003 Achituv & Newman 2002 ogawa et al. 1998 Ross & Newman 2000 Achituv et al. 2009 Achituv & Newman 2002 Achituv & Newman 2002 Ross & Newman 2000 Ross & Newman 2000 ogawa et al. 1998 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 ogawa et al. 1998 Achituv 2001 Ross & Newman 2000 Ross & Newman 2000 ogawa et al. 1998 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Class : Maxillopoda Sub Class : Cirripedia order: Sesslilia Family: balanidae Family: Pyrgomatidae 90 CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa Species Coral taxa Type use Reference Ceratoconcha quarta Cionophorus guillaumae Cionophorus soongi Creusia spinulosa Darwiniella conjugatum Eohoekia chaos Eohoekia nyx Galkinia angustiradiata Galkinia decima Galkinia indica Por Ast Ast Scl Scl Scl Scl Scl Scl Scl o o o o o o o o o o H H H H H H H H H H Scott 1987 Achituv & Newman 2002 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 ogawa et al. 1998 ogawa et al. 1998 Galkinia supraspinulosa Hiroa stubbingsi Hoekia fornix Hoekia monticulariae Hoekia mortenseni Hoekia philippensis Megatrema anglicum Megatrema madreporarium Megatrema oulastreae Neopyrgoma lobata Neotravatha elongatum Nobia conjugatum Nobia grandis Nobia halomitrae Nobia orbicellae Parahoekia aster Pyrgoma cancellatum Pyrgoma japonica Pyrgoma kuri Pyrgoma monticulariae Pyrgoma projectyum Pyrgoma sinica Pyrgopsella annandalei Pyrgopsella stellula Savignium crenatum Savignium elongatum Trevathana dentata Trevathana jensi Trevathana margaretae Trevathana mizrachae Trevathana niuea Trevathana orientale Scl Ast Scl Scl Scl Scl Scl Aga Scl Tub Fav, Goni Scl Scl Scl Scl Scl Den loph Cary Scl Cary Den Scl Scl Goni Scl Goni, lep Fav Fav Plat Gon Fav o o o o o o o o o o o u u o o o o o o o o o o o o u o o o o o o H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H H Ross & Newman 2002 Achituv & Newman 2002 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 Scott 1987 Ross & Newman 2000 Ross & Newman 2002 Ross & Newman 2002 ogawa et al. 1998 ogawa et al. 1998 Ross & Newman 2000 Ross & Newman 2000 Ross & Newman 2000 ogawa et al. 1998 Ross & Newman 2002 Ross & Newman 2002 Ross & Newman 1969 Ross & Newman 2002 Ross & Newman 2002 Ross & Newman 2002 Ross & Newman 2002 Ross & Newman 1973 ogawa et al. 1998 ogawa et al. 1998 brickner et al. 2010 brickner et al. 2010 brickner et al. 2010 ogawa et al. 1998 ogawa et al. 1998 Trevathana paulayi Trevathana sarae Wanella andersonorum Wanella snelliusi Acan Cyph Scl Scl o o o o H H H H ogawa et al. 1998 brickner et al. 2010 Ross & Newman 2000 Ross & Newman 2000 continued 91 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Species Coral taxa Type use Reference Corallonoxia baki Corallonoxia longicauda Corallonoxia mixtibrachium Corallonoxia similis Corallonoxia ventrospinosa Corallovexia brevibrachium Corallovexia dorospina Corallovexia kristenseni Corallovexia longibrachium Corallovexia mediobrachium Pterinopsyllus stirpipes Mea Mea Fav Acr Mont Dip Mont Fav Fav Dip Gal o o o o o o o o o o o F F F F F F F F F F H Stock 1975 Stock 1975 Stock 1975 Stock 1975 Stock 1975 Stock 1975 Stock 1975 Stock 1975 Stock 1975 Stock 1975 Humes 1997a Alteuthellopsis corallina Poc, Acr, Goni, Mer, Plat, Ast Acr Sty, Poc o F Humes 1981 o o H H Humes 1981 Humes 1981 Monp ocu Aga Gon, Gal Gal Gon, Gal Gal Gon, Fung Gon Aga Ech Aga Gon Pav Fung Fung Fung Fung Gon Gal Alv Gal Fung Ser Fung Fung Hydn o o o o o o o o o o o o o o u u u u o o o o o u u o u H F F F F F F F F F F F F F u u u u F F F F F u u F u Humes 1994 Humes 1991a Kim 2006 Humes 1995a Humes 1996a Humes 1995a Humes 1979b Humes & Ho 1968 Humes & Ho 1968 Humes & Stock 1972 Humes 1991a Humes 1992 Humes 1995a Kim 2007 Humes 1996a Kim 2007 Humes 1978a Kim 2003 Humes 1995a Humes 1996a Kim 2003 Humes 1996b Humes 1978a Kim 2003 Humes 1978a Humes 1978a Kim 2007 Sub Class : Copepoda order: Cyclopoida Family: Corallovexiidae Family: Pterinopsyllidae order: Harpacticoida Family: Peltidiidae Family: Tegastidae Tegastes acroporanus Tegastes georgei order: Poecilostomatoida Family: Allopodion mirum Anchimolgidae Anchimolgus abbreviatus Anchimolgus angustus Anchimolgus brevarius Anchimolgus compressus Anchimolgus conformatus Anchimolgus contractus Anchimolgus convexus Anchimolgus digitatus Anchimolgus eparmatoides Anchimolgus exsertus Anchimolgus gibberulus Anchimolgus gigas Anchimolgus gracilipes Anchimolgus gratus Anchimolgus hastatus Anchimolgus latens Anchimolgus maximus Anchimolgus mimeticus Anchimolgus moluccanus Anchimolgus multidentatus Anchimolgus nasutus Anchimolgus notatus Anchimolgus noumensis Anchimolgus orectus Anchimolgus pandus Anchimolgus paragensis 92 CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa Species Coral taxa Type use Reference Anchimolgus partenuipes Anchimolgus prolixipes Anchimolgus punctilis Anchimolgus setellus Anchimolgus tanaus Anchimolgus tenuipes Anchimolgus tridentatus Andrianellus exsertidens Andrianellus papillipes Amarda compta Poc Por Fung Aga ocu Ser Ech Fav Plat Fav u u u o o u u o u o u u u F F u u F u F Kim 2007 Humes & Ho 1968 Humes 1978a Humes 1992 Humes 1991a Kim 2003 Kim 2003 Humes & Stock 1973 Kim 2007 Humes & Stock 1972 Amarda cultrata Amarda curvus Amarda goniastreae Amardopsis merulinae Dumbeana undulatipes Ecphysarion ampullulum Ecphysarion lobophorum Ecphysarion spinulatum Cerioxynus alatus Cerioxynus bandensis Cerioxynus favitocolis Cerioxynus moluccensis Cerioxynus montastreae Cerioxynus oulophylliae Ecphysarion lobophorum Haplomolgus incolumis Haplomolgus montiporae Haplomolgus subdeiciens Humesiella corallicola Kawanolus parangensis Lipochaetes extrusus Odontomolgus actinophorus Odontomolgus bulbalis Odontomolgus campulus Odontomolgus decens Odontomolgus exilipes Odontomolgus lammeus Odontomolgus forhani Odontomolgus fultus Odontomolgus geminus Odontomolgus mucosus Odontomolgus mundulus Fav Goni Goni Mer Psa Acr Acr Acr Fav Fav Fav Fav Fav Fav Acr Monp Monp Monp Hydn Monp Psa Pav Mer Alv Fung Psa Fung Monp Fung Psa Aga Alv o o o o o o u o o o o o o o o o o o o o o o u o o u u o o u o o F F F F F F u F F F F F F F H F H F H H F F u F F u u F F u F F Humes & Stock 1972 Kim 2007 Humes 1985b Humes 1974a Humes 1996b Humes 1993 Humes & Ho 1968 Humes 1993 Humes 1974a Humes 1979c Humes 1974a Humes 1974a Humes 1986 Humes 1986 Humes 1994 Humes 1991b Humes 1994 Humes 1978b Sebastian & Pillai 1973 Humes 1994 Humes 1996b Humes & Frost 1964 Humes 1991a Humes & Ho 1968 Humes 1978a Kim 2003 Kim 2007 Humes 1978b Humes 1978a Kim 2003 Kim 2006 Humes 1974b Odontomolgus parvus Odontomolgus pavonus Odontomolgus pumilis Odontomolgus rhadinus Odontomolgus scitulus Goni Pav Aga Psa Fung u u o o o u u F F F Kim 2007 Kim 2007 Humes 1992 Humes & Ho 1967 Humes 1973 continued 93 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Family: Clausidiidae Family: Rhynchomolgidae Species Coral taxa Odontomolgus unioviger Panjakus bidentis Panjakus directus Panjakus eumeces Panjakus fastigatus Panjakus hydnophorae Panjakus iratus Panjakus necopinus Panjakus parvipes Panjakus platygyrae Aga Poc Fav Hydn Plat Hydn Hydn Fav Plat Plat o o o o o o o o o o F F F F F F F F F F Kim 2006 Kim 2004 Humes 1995b Humes 1991a Kim 2005 Humes & Stock 1973 Kim 2005 Humes 1995b Kim 2005 Humes & Stock 1973 Panjakus saccipes Paraclamocus hiulcus Prionomolgus lanceolatus Rakotoa ceramensis Rakotoa proteus Schedomolgus arcuatipes Schedomolgus dumbensis Schedomolgus exiliculus Schedomolgus idanus Schedomolgus insignellus Schedomolgus majusculus Schedomolgus tener Schedomolgus tenuicaudatus Schedomolgus walteri Scyphuliger aristoides Scyphuliger concavipes Scyphuliger eumorphus Scyphuliger humesi Scyphuliger karangmiensis Scyphuliger latus Scyphuliger longicaudatus Scyphuliger manifestus Scyphuliger paucisurculus Scyphuliger pennauts Scyphuliger pilosis Scyphuliger placidus Scyphuliger tenuatus Scyphuliger vicinus Stockia indica Unicispina latigenitalis Hemicyclops columnaris Hydn Psa Aga Fav Fav Acr Fung Acr Acr Acr Acr Fung Acr lob Acr Acr Acr Acr Acr Acr Acr Acr Acr Acr Acr Acr Acr Acr Fav Acr Por ma. o o o o o o o o o o o o u u o o o o u u u o u u u o o o o o F F F F F F F F F F F F H u u F F F F u u u F u u u F F F F F H Kim 2005 Humes 1997b Humes & Ho 1968 Humes 1979c Humes & Stock 1973 Humes & Ho 1968 Kim 2003 Humes 1993 Humes 1993 Humes 1993 Humes 1993 Humes 1973 Kim 2003 Kim 2003 Humes 1993 Humes 1991a Humes 1993 Kim 2004 Kim 2007 Kim 2003 Kim 2003 Humes 1991a Kim 2003 Kim 2003 Kim 2003 Kim 2004 Humes 1994 Kim 2004 Humes 1994 Humes 1993 Humes 1984a Diallagomolgus productus Diallagomolgus vicinus Isomolgus desmotes Kombia angulata Kombia avitus Kombia curvata Cyph Cyph Ser Psa Por Por ma. o o o o u o F F F F u F Humes 1979d Humes 1979d Dojiri 1988 Humes 1962b Kim 2007 Nair & Pillai 1986 94 Type use Reference CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa Family: Xariiidae Species Coral taxa Type use Reference Kombia imminens Kombia incrassata Mandobius regalis Monomolgus baculigeres Monomolgus psammocorae Monomolgus torulus Monomolgus unihastatus Numboa porosa Pennatulicola corallophilus Ravahina tumida Por ma. Por ma. Pec Por br. Psa Por ma. Por br. Psa Por ma. Por br. o o o o o o o o o o F F F F F F F F F F Humes 1979a Humes 1984b Humes 1991c Humes 1979a Humes & Ho 1967 Humes 1984b Humes & Stock 1973 Humes 1997b Nair & Pillai 1986 Humes & Ho 1968 Spaniomolgus compositus Spaniomolgus crassus Spaniomolgus geminus Wedanus inconstans Xenomolgus varius Hastatus faviae Lipochrus acroporinus Orstomella faviae Orstomella lobophylliae Orstomella yaliuensis Xariia ablusa Xariia acicularis Xariia anomala Xariia anopla Xariia apertipes Xariia basilica Xariia brevicauda Xariia breviramea Xariia bullifera Xariia clavellata Xariia comptula Xariia comata Xariia curtata Xariia decorata Xariia diminuta Xariia dispar Xariia dissona Xariia echinoporae Xariia eminula Xariia exigua Xariia exserens Xariia extensa Ser Sty Sty Gon Por ma. Fav Acr Fav lob Por ma. Acr Aga Acr Monp Cary Acr Alv Acr Acr Aga Hydn Poc Hydn Sty Psa Ech Sty Ech Ser Aga Gal Monp o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F Humes & Frost 1964 Humes & Ho 1968 Humes & Ho 1968 Humes 1978c Humes & Stock 1973 Ho et al. 2010 Humes & Dojiri 1982 Humes & Ho 1968 Humes & Ho 1968 Cheng et al. 2009 Humes & Dojiri 1982 Humes 1985c Humes & Ho 1968 Humes & Dojiri 1982 Humes & Dojiri 1983 Humes 1985c Humes & Ho 1968 Humes 1994 Humes 1985c Humes 1985c Humes & Dojiri 1983 Humes 1962a Humes & Dojiri 1983 Humes & Ho 1968 Humes & Ho 1967 Humes 1962a Humes 1985c Humes & Dojiri 1982 Humes 1985c Humes & Ho 1968 Humes 1985c Humes & Dojiri 1982 Xariia exuta Xariia fastiga Xariia ilata Xariia imbriata Xariia initima Acr Acr Aga Poc Pav o o o o o F F F F F Humes & Dojiri 1982 Humes & Dojiri 1982 Humes 1985c Humes 1985a Humes 1985c continued 95 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Species Coral taxa Xariia issilis Xariia formosa Xariia gerlachi Xariia gibberula Xariia gracilipes Xariia gradata Xariia guttulifera Xariia hadra Xariia hamata Xariia heteromeles Poc Psa Acr Poc Cary Cary Acr Gon Tur Monp o o o o o o o o o o F F F F F F F F F F Humes 1985c Humes 1985c Humes 1994 Humes 1985c Humes & Dojiri 1983 Humes & Dojiri 1983 Humes & Dojiri 1982 Humes & Dojiri 1983 Humes & Ho 1968 Humes 1994 Xariia imitans Xariia imparilis Xariia indica Xariia infrequens Xariia insolita Xariia jugalis Xariia laccadivensis Xariia lacerans Xariia lamellispinosa Xariia levis Xariia linearis Xariia lissa Xariia longa Xariia longicauda Xariia longipes Xariia maldivensis Xariia mediolobata Xariia minax Xariia mucronata Xariia obesa Xariia pectinea Xariia plectrata Xariia quinaria Xariia radians Xariia rasilis Xariia reducta Xariia resex Xariia robusta Xariia rosariae Xariia sabiuraensis Xariia scutipes Xariia sectilis Psa Poc Acr Acr Tub Poc Acr Tur Aga Ser Acr Sty Por br. Acr Pav Poc Alv Cary Acr Poc Acr ocu Poc Alv Acr Ser Gon Acr Acr Acr Acr Poc o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o o F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F F Humes 1985c Humes 1985c Nair 1983 Humes 1962a Cheng et al. 2007 Humes 1985c Nair 1983 Humes 1985c Humes & Ho 1968 Humes 1985c Nair 1983 Humes & Ho 1968 Cheng et al. 2007 Nair 1983 Humes 1962a Humes 1960 Humes & Dojiri 1982 Humes & Dojiri 1983 Humes & Dojiri 1982 Humes & Ho 1968 Humes & Dojiri 1982 Humes 1985c Humes 1985c Humes & Dojiri 1982 Humes 1985c Humes 1962a Humes & Dojiri 1983 Nair 1983 Humes & Dojiri 1982 Humes 1994 Humes & Dojiri 1983 Humes 1985c Xariia serrata Xariia simplex Xariia syntoma Xariia temnura Xariia tenta Xariia tenuis Poc Mer Monp Monp Poc Acr o o o o o o F F F F F F Humes 1962a Humes 1985c Humes 1994 Humes 1994 Humes 1985c Humes 1962a 96 Type use Reference CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa Species Coral taxa Xariia torigera Xariia trituberata Xariia tumorisa Xariia uncinata Xariia umbonata Xariia varilabrata Xariia villosa Zazaranus fungicolis Fav Acr Acr Tur Ser Ser Cyph Fung o o o o o o o o F F F F F F F F Humes 1985c Humes 1994 Humes 1994 Humes 1985c Humes 1985c Humes 1985c Humes & Dojiri 1982 Humes & Dojiri 1983 Fung Fung Monp Gal Gal o o o o o H H H H H Humes 1997a Humes 1997a Humes 1994 Humes 1997a Humes 1997a Barbatia foliata Barbatia fusca Barbatia setigera Por Acr Anac, Monp u u u H H H Zuschin et al. 2001 Mohammed & yassien 2008 Mohammed & yassien 2008 Brachidontes variabilis Fungiacava eilatensis Lithophaga antillarum Sty Fung Cyph, Monp, Goni, Sty, Mont, Dip Ste Sty Mad Por ma. Poc, Acr, Sty Por ma., Ast, Monp, Gon, Cyph Sty Cyph Por, Poc, Fav Goni, Por Goni, Fav Cyph Cyph, Monp Ast, Gon, Fav, Scl Por, Sty Poc Poc u o o H H H Mohammed & yassien 2008 Goreau et al. 1969 Gohar & Soliman 1963 o o o o o o o H H H H H H H Scott 1986 Scott 1988 Soliman 1969 Scott 1986 otter 1937 Wilson 1979 Wilson 1979 o o F o F o o o o u u u H H H H H H H H H H u u Mokady et al. 1991 Soliman 1969 Mohammed & yassien 2008 Scott 1980 Fang & Shen 1988 Wilson 1979 Wilson 1979 Wilson 1979 Gohar & Soliman 1963 Mohammed & yassien 2008 black & Prince 1983 black & Prince 1983 order: Siphonostomatoida Family: Temanus halmaherensis Artotrogidae Tondua tholincola Family: Hetairosyna sororia Asterocheridae Hetairosynopsis bucculentus Madacheres serrulatus Type use Reference Phylum Mollusca Class: bivalvia order: Arcoida Family: Arcidae order: Mytiloida Family: Mytilidae Lithophaga aristata Lithophaga bisulcata Lithophaga cumingiana Lithophaga dixonae Lithophaga hanleyana Lithophaga kuehnelta Lithophaga laevigata Lithophaga lessepsiana Lithophaga lima Lithophaga malaccana Lithophaga nasuta Lithophaga nigra Lithophaga obesa Lithophaga parapurpurea Lithophaga simplex Lithophaga teres Modiolus auriculatus Musculista glaberrima Septifer bilocularis continued 97 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa order: ostreoida Family: ostreidae Family: Pectinidae Family: Spondylidae order: Pterioda Family: Pteriidae order: veneroida Family: veneridae Family: Cardiidae Species Coral taxa Type use Reference Alectryonella plicatula Ostraea sp. Pedum spondyloideum Chlamys madreporarum Chlamys sp. A Chlamys sp. b Mimachlamys lentiginosa Spondylus nicobaricus Spondylus spinosus Por ma. Por ma. Por, Acr Acr Ser Acr Poc Sty Sty F F o u u u u u u H H H u u u u H H Coleman 2003 Coleman 2003 yonge 1967 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 Mohammed & yassien 2008 Mohammed & yassien 2008 Electroma alacorvi Pinctada radiata Acr, Poc Acr, Sty, Monp u u H H Mohammed & yassien 2008 Mohammed & yassien 2008 Gefrarium pectinatum Tridacna crocea Tridacna maxima Sty Por ma. Por, Sty u u u H H H Mohammed & yassien 2008 Hamner & Jones 1976 Mohammed & yassien 2008 Poc Poc u u u u black & Prince 1983 black & Prince 1983 Ser Poc Poc Cary Por ma. u u F F u u u F F H Stella et al. 2010 black & Prince 1983 Glynn et al. 1972 Robertson 1970 Peyrot-Clausade et al. 1992 Por F F Robertson 1970 Poc, Ser Poc Poc Poc Acr Den Tub Den Den Fung Fung Fung Fung Fung Fung Fung u u u u u o o o o o o o o o o o u u u u u F F F F F F F F F F F Stella et al. 2010 black & Prince 1983 black & Prince 1983 black & Prince 1983 Stella et al. 2010 Gittenberger & Gittenberger 2005 Coleman 2003 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Class: Gastropoda order: Caenogastropoda Family: Cerithiidae Cacozeliana granarium Cerithium nesioticum order: littorinimorpha Family: Cypraeidae Family: littorinidae Family: Pediculariidae Family: vermetidae Cypraea asellus Austrolittorina unifasciata Jenneria pustulata Pedicularia decussata Dendropoma maximum order: Neogastropoda Family: Philippia radiata Architectonicidae Family: buccinidae Pisania fasiculata Pisania ignea Family: Pyrene bidentata Collumbellidae Pyrene testudinaria Family: Epitoniidae Cirsotrema sp. A Epidendrium aureum Epidendrium billeeanum Epidendrium dendrophylliae Epidendrium sordidum Epifungium adgranulosa Epifungium adgravis Epifungium adscabra Epifungium hartogi Epifungium hoeksemai Epifungium lochi Epifungium marki 98 CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa Family: Fasciolariidae Family: Mitridae Family: Muricidae Species Coral taxa Epifungium nielsi Epifungium pseudolochi Epifungium pseudotwilae Epifungium twilae Epifungium ulu Epitonium crassicostatum Epitonium graviarmatum Epitonium hartogi Surrepifungium costulatum Surrepifungium ingridae Fung Fung Fung Fung Fung Fung Fung Pler Fung Fung o o o o o o o o o o F F F F F F F F F F Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger & Gittenberger 2005 Gittenberger 2003 Gittenberger et al. 2000 Gittenberger et al. 2000 Surrepifungium oliverioi Surrepifungium patamakanthini Microcolus dunkeri Fung Fung o o F F Gittenberger et al. 2000 Gittenberger & Gittenberger 2005 Poc u u black & Prince 1983 Mitra ferruginea Mitra sp. A Mitra sp. b Coralliophila abreviata Coralliophila australis Coralliophila caribaea Coralliophila costularis Coralliophila erosa Coralliophila imbriata Coralliophila latilirata Coralliophila neritoidea Coralliophila pyriformis Coralliophila violacea Cronia avellana Dicathais textilosa Drupella cornus Poc Poc, Ser, Acr Poc, Ser Acr Scl Acr, Por Acr Acr, Monp Aga Scl Por Tur Por ma. Poc Poc Acr, Monp, Poc, Ser Gal, Monp, Acr Gal, Monp, Acr Acr, Monp Plat Scl Poc Scl Scl Acr Acr Poc Poc, Acr u u u o u o o o u u o o o u u o u u u F F F F F F F F F F u u F Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 brawley & Adey 1982 oliverio 2009 brawley & Adey 1982 Taylor 1978 Robertson 1970 oliverio 2009 oliverio 2009 McClanahan 1997 Coleman 2003 oren et al. 1998 black & Prince 1983 black & Prince 1983 McClanahan 1997 o F boucher 1986 o F Hsieh et al. 2007 o F u o o o u u o o F F F F F F u u F F Morton et al. 2002 Morton & blackmore 2009 yokochi 2004 Glynn et al. 1983 oliverio 2009 oliverio 2009 Stella et al. 2010 Stella et al. 2010 Glynn et al. 1983 Robertson 1970 Drupella fragum Drupella minuta Drupella rugosa Ergalatax margariticola Habromorula spinosa Latiaxis hindsii Magilus antiquus Magilus lamarkii Morula sp. A Morula sp. b Muricopsis zeteki Quoyula madreporarum Type use Reference continued 99 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Family: Phasianellidae Family: Strombidae Family: Triviidae Species Coral taxa Quoyula monodonta Rapa incurvus Phasianella sp. A Poc, Por br. Scl Acr o u u F F u Robertson 1970 oliverio 2009 Stella et al. 2010 Strombus mutabilis Strombus sp. A Strombus sp. b Strombus sp. C Trivia merces Trivia oryza Ser Poc, Ser Acr Acr Poc Poc, Ser, Acr u u u u u u u u u u u u Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 black & Prince 1983 Stella et al. 2010 Poc u u black & Prince 1983 Poc Poc Poc Poc Poc Poc Poc Ser Poc, Acr Acr Acr u u u u u u u u u u u u u u u u u u u u u u black & Prince 1983 black & Prince 1983 black & Prince 1983 black & Prince 1983 black & Prince 1983 black & Prince 1983 black & Prince 1983 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 Phestilla melanobranchia Phestilla sibogae Phestilla lugubris Phestilla minor Den, Tub Por, Gon, Tub Por, Gon, Tub Por, Gon, Tub o o o o F F F F Harris 1975 Harris 1975 Rudman 1981 Rudman 1981 Cryptoplax sp. Poc, Ser, Acr u u Stella et al. 2010 Coscinasterias calamaria Poc u u black & Prince 1983 Acanthaster planci Acr, Poc, Mus, Fav, Por Por ma. Por ma. o F Pearson & Endean 1969 F F F F Thomassin 1976 Thomassin 1976 order: vetigastropoda Family: Scutus antipodes Fissurellidae Family: Haliotidae Haliotis varia Family: Trochidae Clanculus denticulatus Clanculus personatus Clanculus plebejus Herpetopoma aspersa Family: Turbinidae Australium tentorium Turbo argyrostomus Turbo brunneus Turbo sp. A Turbo sp. b Turbo sp. C Type use Reference Class: Nudibranchia order: Aeolidina Family: Tergipedidae Class: Polyplacophora order: Chitonida Family: Cryptoplacidae Phylum Echinodermata Class: Asteroidea order: Forcipulatida Family: Asteriidae order: Spinulosida Family: Acanthasteridae Family: Echinasteridae Echinaster luzonicus Echinaster purpureus 100 CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa order: valvatida Family: Asterinidae Family: ophidiasteridae Family: oreasteridae Species Coral taxa Type use Reference Asterina anomala Asterina atyphoida Asterina sp. Linckia laevigata Nardoa variolata Pharia pyramidata Culcita novaeguineae Culcita schmideliana Nidorellia armata Pentaceraster cumingi Acr Poc Acr Por ma. Por ma. Poc Poc Gal, Gon Pav Psa F u F F F F F F F F F u F F F F F F F F yamaguchi 1975 black & Prince 1983 yamaguchi 1975 Thomassin 1976 Thomassin 1976 Dana & Wolfson 1970 Glynn & Krupps 1986 Thomassin 1976 Glynn et al. 1983 Glynn 2004 Eucidaris metularia Eucidaris thouarsii Ser Pav, Poc u F u S Stella et al. 2010 Glynn et al. 1983 Colobocentrotus atratus Echinometra lucunter Echinometra mathaei Echinometra viridis Ser, Acr Scl Scl Scl u F F F u S S S Stella et al. 2010 ogden 1977 Herring 1972 Grifin et al. 2003 F F S S Herring 1972 Glynn 1988 Diadema mexicanum Diadema savignyi Diadema setosum Echinothrix calamaris Echinothrix diadema Scl Acr, Aga, Mad, Mont, Por Scl Scl Scl Scl Scl F F F F F S S S S S Glynn 1988 bak 1990 Herring 1972 Herring 1972 bak 1990 Echinoneus cyclostomus Scl F S Herring 1972 Scl F S Herring 1972 Poc u u black & Prince 1983 Scl Scl F F S S Herring 1972 Herring 1972 Class: Echinoidea order: Cidaroida Family: Cidaridae order: Echinoida Family: Echinometridae order: Echinothuroida Family: Astropyga radiata Diadematidae Diadema antillarum order: Holectypoida Family: Echinoneidae order: Phymosomatoida Family: Stomopneustes variolaris Stomopneustidae order: Temnopleuroida Family: Temnopleurus michaelseni Temnopleuridae Family: Microcyphus rousseaui Toxopneustidae Tripneustes gratilla continued 101 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Species Coral taxa Type use Reference Amphipholis squamata Amphiura luetkeni Ophiactis savignyi Ophiactis sp. A Clarkcoma canaliculata Ophiocoma dentata Ophiocoma erinaceus Ophiocoma sp. A Ophiocoma sp. b Ophiocoma sp. C Ophiocoma sp. D Ophiocoma sp. E Ophiocoma occidentalis Ophiocomella sexradia Macrophiothrix sp. Ophiomastix sp. Ophiothrix acestra Ophiothrix sp. A Ophiothrix sp. b Poc Poc Poc Acr Poc Poc Poc, Ser, Acr Poc, Ser, Acr Poc, Ser, Acr Poc, Ser Ser, Acr Poc Poc Poc Ser, Acr Poc, Ser Poc Poc, Ser, Acr Poc, Ser, Acr u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u u Austin et al. 1980 Austin et al. 1980 Austin et al. 1980 Stella et al. 2010 black & Prince 1983 black & Prince 1983 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 Stella et al. 2010 black & Prince 1983 Austin et al. 1980 Stella et al. 2010 Stella et al. 2010 Austin et al. 1980 Stella et al. 2010 Stella et al. 2010 Chloeia lava Chloeia fusca Eurythoe complanata Hermodice carunculata Notopygos crinita Scl Scl Mont Por Mont F F F F F u u u F u Fossa & Nilsen 2000 Fossa & Nilsen 2000 Fossa & Nilsen 2000 Marsden 1962 Ebbs 1966 Pseudovermilia madracicola Spirobranchus corniculatus Poc Por ma., Por br., Acr, Aga, Poc, Mont Por ma., Pav Por ma. Por ma. Por ma. Por ma. Por ma. Por ma. o o u H ten Hove 1994 ten Hove 1994 o u o u o u o H H H H H H H ten Hove 1994 Fossa & Nilsen 2000 Devantier et al. 1986 Fossa & Nilsen 2000 ten Hove 1994 Fossa & Nilsen 2000 ten Hove 1994 Class: ophiuroidea order: ophiurida Family: Amphiuridae Family: ophiactidae Family: ophiocomidae Family: ophiotrichidae Phylum Annelida Class: Polychaeta order: Amphinomida Family: Amphinomidae order: Canalipalpata Family: Serpulidae Spirobranchus gardineri Spirobranchus gaymardi Spirobranchus giganteus Spirobranchus incrassatus Spirobranchus nigranucha Spirobranchus paumotanus Spirobranchus polycerus 102 CoRAl-ASSoCIATED INvERTEbRATES Higher Taxa order: Eunicida Family: Eunicidae Family: lumbrineridae Family: oenonidae order: Phyllodocida Family: Aphroditidae Family: Polynoidae order: Spionida Family: Chaetopteridae Family: Spionidae Species Coral taxa Type use Reference Spirobranchus spinosus Spirobranchus tetraceros Por ma. Por ma. Pav u o H H Fossa & Nilsen 2000 ten Hove 1994 Eunice loridana Eunice mutilata Eunice pennata Eunice schemacephala Lumbrineris labellicola ocu Mont Mad Mont Den, Cary, Fla F F F F o u H u H u britayev 1981 Ebbs 1966 Fauchald 1992 Ebbs 1966 Zibrowius et al. 1975 Oenone fulgida Mont F u Ebbs 1966 Pontogenia sericoma Mont F u Ebbs 1966 Harmothoe aculeata Hermenia verruculosa Hololepidella nigropunctata Lepidonotus variabilis Mont Mont Fung Mont F F o F u u u u Ebbs 1966 Ebbs 1966 Pettibone 1993 Ebbs 1966 Spiochaetopterus sp. Monp o H bergsma 2009 Dipolydora armata lep F H okuda 1937 Cliona laticavicola Cliona mucronata Cliona pocillopora Acr Poc Poc o o o S S S Pang 1973 bautista-Guerrero et al. 2006 bautista-Guerrero et al. 2006 Callyspongia californica Haliclona caerulea Chalinula nematifera Amphimedon texotli Aka cryptica Poc Poc Poc Poc Poc F F o F u S S S S S Cruz-barraza & Carballo 2008 Cruz-barraza & Carballo 2008 Cruz-barraza & Carballo 2008 Cruz-barraza & Carballo 2008 Carballo et al. 2007 Poc Mont, Aga, Por ma., Mus Poc F F S S Cruz-barraza & Carballo 2008 Goreau & Hartman 1966 F S Cruz-barraza & Carballo 2008 Phylum Porifera Class: Demospongiae order: Hadromerida Family: Clionaidae order: Haplosclerida Family: Callyspongiidae Family: Chalinidae Family: Niphatidae Family: Phloeodictyidae order: Poecilosclerida Family: Mycalidae Mycale cecilia Mycale laevis Mycale magnirhaphidifera continued 103 JESSICA S. STEllA, MoRGAN S. PRATCHETT, PAT A. HuTCHINGS & GEoFFREy P. JoNES Higher Taxa Species Coral taxa Type use Reference Phylum Platyhelminthes Class: Turbellaria order: Acoela Family: Convolutidae Family: Sagittiferidae order: Polycladida Family: Prosthiostomidae Waminoa brickneri Waminoa litus Convolutriloba hastifera Convolutriloba longiissura Convolutriloba macropyga Convolutriloba retrogemma Haplodiscus sp. Notoplana tremellaris Fav Por ma Goni, Plat, Ple Ple Scl Ple Por Ple u u u u u u u u F F F F F F F F ogunlana et al. 2005 ogunlana et al. 2005 Winsor 1990 bartolomaeus & balzer 1997 Shannon & Achatz 2007 Winsor 1990 Trench & Winsor 1987 Delbeek & Sprung 1997 Prosthiostomum sp. Monp o F Jokiel & Townsley 1974 Por ma. Cary, Den u F H H Peyrot-Clausade et al. 1992 Hoeksema & best 1991 loph u u Dilly & Ryland 1985 Phylum Sipuncula Class: Phascolosomatidea order: Aspidosiphonida Family: Aspidosiphon elegans Aspidosiphonidae Aspidosiphon muelleri Phylum Hemichordata Class: Pterobranchia order: Rhabdopleuroidea Family: Rhabdopleura normani Rhabdopleuridae Key to abbreviations Type: o = obligate symbiont, F = Facultative symbiont, u = unknown. use: F = Food, H = Habitat, S = Substratum, u = unknown Coral taxa: Acan = Acanthastraea, Acr = Acropora, Aga = Agariciidae, Anac = Anacropora, Ast = Astreopora, Cary = Caryophyllidae, Cyph = Cyphastrea, Den = Dendrophyllidae, Dip = Diplora, Ech = Echinopora, Eup = Euphyllia, Fav = Favia + Favites, Fla = Flabellidae, Fung = Fungiidae, Gal = Galaxea, Gon = Goniopora, Goni = Goniastrea, Hydn = Hydnophora, lep = Leptastrea, lob = Lobophyllia, loph = Lophelia, Mad = Madracis, Mea = Meandrina, Mer = Merulina, Monp = Montipora, Mont = Montastrea, Mus = Mussidae, ocu = Oculina, Pach = Pachyseris, Pav = Pavona, Pec = Pectiniidae, Pla t = Platygyra, Pler = Plerogyra, Ple = Plesiastrea, Psa = Psammocora, Por = Porites (ma, massive, br, branching), Poc = Pocillopora, Rhi = Rhizangiidae, Ser = Seriatopora, Sid = Siderastrea, Ste = Stephanocoenia, Sty = Stylophora, Scl = unknown scleractinian coral, Tham = Thamnastrea, Tub = Tubastrea, Tur = Turbinaria 104

RELATED PAPERS

RELATED TOPICS

Log In

Coral-associated invertebrates: diversity, ecological importance and vulnerability to disturbance

Coral-associated invertebrates: diversity, ecological importance and vulnerability to disturbance

Related Papers

RELATED PAPERS

RELATED TOPICS