Implications of reef ecosystem change for the stability and maintenance of coral reef islands

Coral reef islands are among the most vulnerable environments on Earth to climate change because they are low lying and largely constructed from unconsolidated sediments that can be readily reworked by waves and currents. These sediments derive entirely from surrounding coral reef and reef flat environments and are thus highly sensitive to ecological transitions that may modify reef community composition and productivity. How such modifications – driven by anthropogenic disturbances and on‐going and projected climatic and environmental change – will impact reef island sediment supply and geomorphic stability remains a critical but poorly resolved question. Here, we review the unique ecological–geomorphological linkages that underpin this question and, using different scenarios of environmental change for which reef sediment production responses can be projected, explore the likely resilience of different island types. In general, sand‐dominated islands are likely to be less resilient than those dominated by rubble grade material. However, because different islands typically have different dominant sediment constituents (usually either coral, benthic foraminifera or Halimeda) and because these respond differently to individual ecological disturbances, island resilience is likely to be highly variable. Islands composed of coral sands are likely to undergo major morphological change under most near‐future ecological change scenarios, while those dominated by Halimeda may be more resilient. Islands composed predominantly of benthic foraminifera (a common state through the Pacific region) are likely to exhibit varying degrees of resilience depending upon the precise combination of ecological disturbances faced. The study demonstrates the critical need for further research bridging the ecological–geomorphological divide to understand: (1) sediment production responses to different ecological and environmental change scenarios; and (2) dependant landform vulnerability.     

Growth of reef fishes in response to live coral cover

Although the global decline in coral reef health is likely to have profound effects on reef associated fishes, these effects are poorly understood. While declining coral cover can reduce the abundance of reef fishes through direct effects on recruitment and/or mortality, recent evidence suggests that individuals may survive in disturbed habitats, but may experience sublethal reductions in their condition. This study examined the response of 2 coral associated damselfishes (Pomacentridae), Chrysiptera parasema and Dascyllus melanurus, to varying levels of live coral cover. Growth, persistence, and the condition of individuals were quantified on replicate coral colonies in 3 coral treatments: 100% live coral (control), 50% live coral (partial) and 0% live coral (dead). The growth rates of both species were directly related to the percentage live coral cover, with individuals associated with dead corals exhibiting the slowest growth, and highest growth on control corals. Such differences in individual growth between treatments were apparent after 29 d. There was no significant difference in the numbers of fishes persisting or the physiological condition of individuals between different treatments on this time-scale. Slower growth in disturbed habitats will delay the onset of maturity, reduce lifetime fecundity and increase individual's vulnerability to gape-limited predation. Hence, immediate effects on recruitment and survival may underestimate the longer-term impacts of declining coral on the structure and diversity of coral-associated reef fish communities.     

Caribbean mesophotic coral ecosystems are unlikely climate change refugia

Deeper coral reefs experience reduced temperatures and light and are often shielded from localized anthropogenic stressors such as pollution and fishing. The deep reef refugia hypothesis posits that light‐dependent stony coral species at deeper depths are buffered from thermal stress and will avoid bleaching‐related mass mortalities caused by increasing sea surface temperatures under climate change. This hypothesis has not been tested because data collection on deeper coral reefs is difficult. Here we show that deeper (mesophotic) reefs, 30–75 m depth, in the Caribbean are not refugia because they have lower bleaching threshold temperatures than shallow reefs. Over two thermal stress events, mesophotic reef bleaching was driven by a bleaching threshold that declines 0.26 °C every +10 m depth. Thus, the main premise of the deep reef refugia hypothesis that cooler environments are protective is incorrect; any increase in temperatures above the local mean warmest conditions can lead to thermal stress and bleaching. Thus, relatively cooler temperatures can no longer be considered a de facto refugium for corals and it is likely that many deeper coral reefs are as vulnerable to climate change as shallow water reefs.     

珊瑚礁生态保护与管理研究

珊瑚礁以其极高的生物多样性和生物生产力以及优美的自然景观 ,为人们提供了生活需要和游乐的资源 ,但同时也受到过度利用的破坏 ,尤其是近年来显得更为严重 ,因而珊瑚礁的生态保护与管理成为近 2 0年来倍受关注的问题。本文回顾了国内外珊瑚礁生态保护和管理的一些研究成果 ,通过自然和社会经济调查 ,并根据保护、研究和可持续利用的原则 ,将雷州半岛灯楼角珊瑚礁保护区划分为野生区 ,保护区、季节性封闭区和一般使用区 ,并强调公众参与、社区组织和领导组成、教育和培训、资源管理等为保护和管理中的措施     

Analysis on variation trend of coral reef in Xisha

Xisha Islands, located in the northern part of South China Sea, consist of more than 20 islands and atolls. The coral reef of Xisha Islands belongs to the typical ocean distribution of world’s coral reefs, its ecosystem is the most typical in our country and hermatypic coral species are about three-quarters of the total coral species in China. It is addressed with the oldest coral reef community which of the most original and valuable in China. The previous research shows that the islands studded in South China Sea such as Xisha Islands have important influence on the formation of coral reefs along the mainland coast by multiplying and migrating from south to the north. It is supplementary sources of coastal coral larvae in Hainan and Guangdong. Therefore, carrying out the monitoring of coral reef community ecosystem is of great significance to the ecological protection. By the Manta tow and the Line Intercept Transect method, five stations (Xisha Yong xing dao, Shi dao, Xisha zhou, Zhao Shu dao and Bei dao) on Xisha ecological monitoring area were monitored continually from 2005 to 2009. We compared the index changes of Hermatypic coral’s species, coverage and recruitment, and combining with Condition Index, Succession Index, and the variation trend of hard coral in Xisha were analyzed.
The results show that, from 2005 to 2009, the coverage of living hermatypic corals in ecological monitoring area is sharply reduced from 68.19% to 7.93%, while the dead coral coverage is sharply increased from 4.70% to 72.90%. Coral recruitment is reduced from 1.18 ind/100 m² to 0.07 ind/100 m², hermatypic coral species decreased from 87 to 35. In 2005, the health of coral reefs was very good, and the Condition Index was 1.097. However, the Condition Index cut down to a very low degree in 2009. It was only ?0.880. The Succession Index belonged to “very low degree” from 2005 to 2009, and the numerical value was gradually reduce from ?0.984 to ?1.876.
As a whole, hermatypic corals are serious degrade regionally and caused great change to the coral structure and biodiversity, this will lead to a continuous degradation of coral reefs.     

Analysis on variation trend of coral reef in Xisha

Xisha Islands, located in the northern part of South China Sea, consist of more than 20 islands and atolls. The coral reef of Xisha Islands belongs to the typical ocean distribution of world’s coral reefs, its ecosystem is the most typical in our country and hermatypic coral species are about three-quarters of the total coral species in China. It is addressed with the oldest coral reef community which of the most original and valuable in China. The previous research shows that the islands studded in South China Sea such as Xisha Islands have important influence on the formation of coral reefs along the mainland coast by multiplying and migrating from south to the north. It is supplementary sources of coastal coral larvae in Hainan and Guangdong. Therefore, carrying out the monitoring of coral reef community ecosystem is of great significance to the ecological protection. By the Manta tow and the Line Intercept Transect method, five stations (Xisha Yong xing dao, Shi dao, Xisha zhou, Zhao Shu dao and Bei dao) on Xisha ecological monitoring area were monitored continually from 2005 to 2009. We compared the index changes of Hermatypic coral’s species, coverage and recruitment, and combining with Condition Index, Succession Index, and the variation trend of hard coral in Xisha were analyzed.
The results show that, from 2005 to 2009, the coverage of living hermatypic corals in ecological monitoring area is sharply reduced from 68.19% to 7.93%, while the dead coral coverage is sharply increased from 4.70% to 72.90%. Coral recruitment is reduced from 1.18 ind/100 m² to 0.07 ind/100 m², hermatypic coral species decreased from 87 to 35. In 2005, the health of coral reefs was very good, and the Condition Index was 1.097. However, the Condition Index cut down to a very low degree in 2009. It was only ?0.880. The Succession Index belonged to “very low degree” from 2005 to 2009, and the numerical value was gradually reduce from ?0.984 to ?1.876.
As a whole, hermatypic corals are serious degrade regionally and caused great change to the coral structure and biodiversity, this will lead to a continuous degradation of coral reefs.     

Habitat choice,recruitment and the response of coral reef fishes to coral degradation

The global degradation of coral reefs is having profound effects on the structure and species richness of associated reef fish assemblages. Historically, variation in the composition of fish communities has largely been attributed to factors affecting settlement of reef fish larvae. However, the mechanisms that determine how fish settlers respond to different stages of coral stress and the extent of coral loss on fish settlement are poorly understood. Here, we examined the effects of habitat degradation on fish settlement using a two-stage experimental approach. First, we employed laboratory choice experiments to test how settlers responded to early and terminal stages of coral degradation. We then quantified the settlement response of the whole reef fish assemblage in a field perturbation experiment. The laboratory choice experiments tested how juveniles from nine common Indo-Pacific fishes chose among live colonies, partially degraded colonies, and dead colonies with recent algal growth. Many species did not distinguish between live and partially degraded colonies, suggesting settlement patterns are resilient to the early stages of declining coral health. Several species preferred live or degraded corals, and none preferred to associate with dead, algal-covered colonies. In the field experiment, fish recruitment to coral colonies was monitored before and after the introduction of a coral predator (the crown-of-thorns starfish) and compared with undisturbed control colonies. Starfish reduced live coral cover by 95–100%, causing persistent negative effects on the recruitment of coral-associated fishes. Rapid reductions in new recruit abundance, greater numbers of unoccupied colonies and a shift in the recruit community structure from one dominated by coral-associated fishes before degradation to one predominantly composed of algal-associated fish species were observed. Our results suggest that while resistant to coral stress, coral death alters the process of replenishment of coral reef fish communities.     

Coral reef encruster communities and carbonate production in cryptic and exposed coral reef habitats along a gradient of terrestrial disturbance

Encrusting calcareous organisms such as bryozoans, crustose coralline algae (CCA), foraminiferans, and serpulid worms are integral components of tropical framework-building reefs. They can contribute calcium carbonate to the reef framework, stabilise the substrate, and promote larval recruitment of other framework-building species (e.g. coral recruits). The percentage cover of encrusting organisms and their rates of carbonate production (g m⁻² year⁻¹) were assessed at four sites within a coastal embayment, along a gradient of riverine influence (high-low). As the orientation and type of substrate is thought to influence recruitment of encrusting organisms, organisms recruiting to both natural (the underside of platy corals) and experimental substrates were assessed. The effect of substrate exposure under different levels of riverine influence was assessed by orientating experimental substrates to mimic cryptic and exposed reef habitats (downwards-facing vs upwards-facing tiles) at each site. Cryptic experimental tiles supported similar encruster assemblages to those recruiting to the underneath (cryptic side) of platy corals, suggesting that tiles can be used as an experimental substrate to assess encruster recruitment in reef systems. Encruster cover, in particular CCA, and carbonate production was significantly higher at low-impact (clear water), high wave energy sites when compared to highly riverine impacted (turbid water), low wave energy sites. Cryptically orientated substrates supported a greater diversity of encrusting organisms, in particular serpulid worms and bryozoans. The inverse relationships observed between riverine inputs and encrusters (total encruster cover and carbonate production) have implications for both the current and future rates and styles of reefal framework production.     

The area of potential shallow-water tropical and subtropical coral ecosystems in the United States

Geographic information system-based analysis was used to derive comprehensive, consistent estimates of the potential area of broadly defined, shallow-water, tropical and subtropical coral ecosystems within the territorial sea and exclusive economic zone of the United States. A coral ecosystem is composed of habitats including unconsolidated sediment, mangrove, hermatypic coral, colonized hardbottom, and submerged vegetation, and major structural zones like reef crest, lagoon, and fore reef. This broad definition reflects the importance of both reef and non-reef habitats and structural zones in the function of these ecosystems. Nautical charts, published by the National Oceanic and Atmospheric Administration’s Office of the Coast Survey, provide a consistent source of 10-fathom (∼18 m) and 100-fathom (∼183 m) depth curve information. The 10-fathom or 100-fathom depth curves are used as surrogates for the potential distribution and extent of shallow-water coral ecosystems in tropical and subtropical U.S. waters. An estimated 36,813 sq·km area has been identified where coral ecosystems can potentially be found in waters less than 10 fathoms (18 m) deep. In addition, an estimated 143,059 sq·km area has been identified where coral ecosystems potentially can be found in U.S. waters at depths down to 100 fathoms (183 m). Results also indicate that previous studies underestimated the extent of potential coral ecosystems for some locations in U.S. tropical and subtropical waters by as much as 100% and that the regional distribution of coral ecosystems has been incorrectly reported.     

Characterization of archaeal symbionts of sponges from the coral reef ecosystems of the Gulf of Mannar,Southeast coast of India

Sponges accommodate a diverse group of microorganisms with varied metabolic capabilities. The bacterial associates of sponges are widely studied while our understanding of archaeal counterparts is scanty. In the present study, we report the archaeal associates of two sponges, Pseudoceratina purpurea (NCBI barcode: KX454492) and Cinachyra sp. (NCBI barcode: KX454495), found in the coral reef ecosystems of Gulf of Mannar, India. Archaea in the water column was predominated by members of class Halobacteria of Phylum Euryarchaeota (97%) followed by a minor fraction (3%) of Nitrosopumilus sp. of phylum Thaumarchaeota. Interestingly, Thaumarchaeota was identified as the sole archaeal population associated with the two sponges studied, among which Nitrosopumilus sp. occuppied 80 and 100% of the sequences in the clone library of P. purpurea and Cinachyra sp. respectively. Other archaea found in the P. purpurea were Nitrososphaera (10%) and unclassified ones (10%). The study identified Nitrosopumilus sp. as a unique symbiotic archaeon of sponges, P. purpurea and Cinachyra sp. The existence of host driven factors in selecting specific associates from a diverse group of archaea in the environment may need further investigations.     

Monitoring coral bleaching using a colour reference card

Assessment of the extent of coral bleaching has become an important part of studies that aim to understand the condition of coral reefs. In this study a reference card that uses differences in coral colour was developed as an inexpensive, rapid and non-invasive method for the assessment of bleaching. The card uses a 6 point brightness/saturation scale within four colour hues to record changes in bleaching state. Changes on the scale of 2 units or more reflect a change in symbiont density and chlorophyll a content, and therefore the bleaching state of the coral. When used by non-specialist observers in the field (here on an intertidal reef flat), there was an inter-observer error of ± 1 colour score. This technique improves on existing subjective assessment of bleaching state by visual observation and offers the potential for rapid, wide-area assessment of changing coral condition.     

Foraminiferal assemblage and reef check census in coral reef health monitoring of East Brazilian margin

Human activity is changing environmental conditions on a global scale. Among the ecosystems that are affected by human activities, coral reefs are among the most prominent. In Brazil, the coral reefs of the Corumbau Marine Extractive Reserve (CMER) and Abrolhos National Marine Park (ANMP) in Bahia state have some of the highest coral cover in the South Atlantic Ocean. Hard coral cover, algal cover, and foraminiferal population distribution patterns were used to assess the coral reef benthic environments, and define a background that can be used in worldwide comparisons in future studies. To compare these two monitoring approaches in different coral reef environments, relative frequency data for occurrence of hard coral and algal cover, using point-intercept transects as proposed by the Reef Check protocol, and foraminiferal samples were collected from Corumbau (nearshore) and Abrolhos (offshore) in April 2005. The foraminiferal assemblage was evaluated using the FORAM index (FI — Foraminifera in Reef Assessment and Monitoring), which provides a numeric diagnosis of suitability of benthic habitat to support calcifying organisms that host algal symbionts, originally developed for Caribbean reef areas. Coral cover in the surveyed areas, both in Corumbau and in Abrolhos, ranged from 13% to 37%, while high foraminiferal diversities (H') were found in all stations. Dominance of symbiont-bearing taxa of Amphistegina lessonii and Archaias angulatus only occurred at two shallow stations, Mato Verde and Siriba, both in Abrolhos, where FI > 4.00. Stations located in Corumbau and Abrolhos had FI values < 4.00. Q-mode cluster analysis showed that foraminifers have specific preferences for physical conditions, especially hydrodynamics and light availability, which influence the FI index. Although coral cover in these areas can be considered good by regional standards, foraminifer analysis showed that the benthic system was unfavorable for symbiont-bearing foraminiferal species at most stations. This discrepancy reveals that the FI must be used with caution in areas other than the northwestern Atlantic and Caribbean where it was developed, and that some coral species can thrive in muddier conditions than can most symbiont-bearing foraminifers.     

Future habitat suitability for coral reef ecosystems under global warming and ocean acidification

Rising atmospheric CO₂ concentrations are placing spatially divergent stresses on the world's tropical coral reefs through increasing ocean surface temperatures and ocean acidification. We show how these two stressors combine to alter the global habitat suitability for shallow coral reef ecosystems, using statistical Bioclimatic Envelope Models rather than basing projections on any a priori assumptions of physiological tolerances or fixed thresholds. We apply two different modeling approaches (Maximum Entropy and Boosted Regression Trees) with two levels of complexity (one a simplified and reduced environmental variable version of the other). Our models project a marked temperature‐driven decline in habitat suitability for many of the most significant and bio‐diverse tropical coral regions, particularly in the central Indo‐Pacific. This is accompanied by a temperature‐driven poleward range expansion of favorable conditions accelerating up to 40–70 km per decade by 2070. We find that ocean acidification is less influential for determining future habitat suitability than warming, and its deleterious effects are centered evenly in both hemispheres between 5° and 20° latitude. Contrary to expectations, the combined impact of ocean surface temperature rise and acidification leads to little, if any, degradation in future habitat suitability across much of the Atlantic and areas currently considered ‘marginal’ for tropical corals, such as the eastern Equatorial Pacific. These results are consistent with fossil evidence of range expansions during past warm periods. In addition, the simplified models are particularly sensitive to short‐term temperature variations and their projections correlate well with reported locations of bleaching events. Our approach offers new insights into the relative impact of two global environmental pressures associated with rising atmospheric CO₂ on potential future habitats, but greater understanding of past and current controls on coral reef ecosystems is essential to their conservation and management under a changing climate.     

Ecological structure of assemblages of coral reef fishes on isolated patch reefs

A 9-year study of the structure of assemblages of fish on 20 coral patch reefs, based on 20 non-manipulative censuses, revealed a total of 141 species from 34 families, although 40 species accounted for over 95% of sightings of fish. The average patch reef was 8.5 m² in surface area, and supported 125 fish of 20 species at a census. All reefs showed at least a two-fold variation among censuses in total numbers of fish present, and 12 showed ten-fold variations. There was also substantial variation in the composition and relative abundances of species present on each patch reef, such that censuses of a single patch reef were on average about 50% different from each other in percent similarity of species composition (Czekanowski's index). Species differed substantially in the degree to which their numbers varied from census to census, and in the degree to which their dispersion among patch reefs was modified from census to census. We characterize the 40 most common species with respect to these attributes. The variations in assemblage structure cannot be attributed to responses of fish to a changing physical structure of patch reefs, nor to the comings and goings of numerous rare species. Our results support and extend earlier reports on this study, which have stressed the lack of persistant structure for assemblages on these patch reefs. While reef fishes clearly have microhabitat preferences which are expressed at settlement, the variations in microhabitat offered by the patch reefs are insufficient to segregate many species of fish by patch reef. Instead, at the scale of single patch reefs, and, to a degree, at the larger scale of the 20 patch reefs, most of the 141 species of fish are distributed without regard to differences in habitat structure among reefs, and patterns of distribution change over time. Implications for general understanding of assemblage dynamics for fish over more extensive patches of reef habitat are considered.     

Grazing pressure of herbivorous coral reef fishes on low coral-cover reefs

The impact of grazing by herbivorous fishes (Acanthuridae, Scaridae, and Pomacentridae) on low coral-cover reefs was assessed by measuring rates of benthic algal production and consumption on inshore and offshore reefs in the upper Florida Keys. Algal production rates, determined in situ with caged and uncaged experimental plates, were low (mean 1.05 g C m⁻² day⁻¹) and similar among reef types. Algal consumption rates were estimated using two different models, a detailed model incorporating fish bite rates and algal yield-per-bite for one species extrapolated to a guild-wide value, and a general regression relating fish biomass to algal consumption. Algal consumption differed among reef types: a majority of algal production was consumed on offshore reefs (55–100%), whereas consumption on inshore patch reefs was 31–51%. Spatial variation in algal consumption was driven by differences in herbivorous fish species composition, density, and size-structure among reef types. Algal consumption rates also varied temporally due to seasonal declines in bite rates and intermittent presence of large-bodied, vagile, schooling species. Spatial coherence of benthic community structure and temporal stability of algal turf over 3 years suggests that grazing intensity is currently sufficient to limit further spread of macroalgal cover on these low coral-cover reefs, but not to exclude it from the system.     

Extinction vulnerability of coral reef fishes

With rapidly increasing rates of contemporary extinction, predicting extinction vulnerability and identifying how multiple stressors drive non-random species loss have become key challenges in ecology. These assessments are crucial for avoiding the loss of key functional groups that sustain ecosystem processes and services. We developed a novel predictive framework of species extinction vulnerability and applied it to coral reef fishes. Although relatively few coral reef fishes are at risk of global extinction from climate disturbances, a negative convex relationship between fish species locally vulnerable to climate change vs. fisheries exploitation indicates that the entire community is vulnerable on the many reefs where both stressors co-occur. Fishes involved in maintaining key ecosystem functions are more at risk from fishing than climate disturbances. This finding is encouraging as local and regional commitment to fisheries management action can maintain reef ecosystem functions pending progress towards the more complex global problem of stabilizing the climate.     

Parasites of recruiting coral reef fish larvae in New Caledonia

Recruiting coral reef fish larvae from 38 species and 19 families from New Caledonia were examined for parasites. We found 13 parasite species (Platyhelminthes: Monogenea, Cestoda and Trematoda) but no acanthocephalan, crustacean or nematode parasites. Over 23% of individual fish were infected. Didymozoid metacercariae were the most abundant parasites. We conclude that most of the parasites are pelagic species that become ‘lost’ once the fish larvae have recruited to the reef. Larval coral reef fish probably contribute little to the dispersal of the parasites of the adult fish so that parasite dispersal is more difficult than that of the fish themselves.     

Impact of micropredatory gnathiid isopods on young coral reef fishes

The ecological role of parasites in the early life-history stages of coral reef fish, and whether this varies between fish with and without a pelagic phase, was investigated. The susceptibility to, and effect of reef-based micropredatory gnathiid isopods on larval, recently settled, and juvenile fishes was tested using two damselfishes (Pomacentridae): Neopomacentrus azysron, which has pelagic larvae, and Acanthochromis polyacanthus, which does not. When larval and recently settled stages of N. azysron and very young A. polyacanthus juveniles (smaller than larval N. azysron) were exposed to one or three gnathiids, the proportion of infections did not vary significantly among the three host types or between the number of gnathiids to which the fish were exposed. The overall infection was 35%. Mortality, however, differed among the three gnathiid-exposed host types with most deaths occurring in larval N. azysron; no mortalities occurred for recently settled N. azysron exposed to one or three gnathiids, and A. polyacanthus exposed to one gnathiid. Mortality did not differ significantly between larval N. azysron and A. polyacanthus juveniles, failing to provide support for the hypothesis that reef-based A. polyacanthus juveniles are better adapted to gnathiid attack than fish with a pelagic phase. The study suggests that settling on the reef exposes young fish to potentially deadly micropredators. This supports the idea that the pelagic phase may allow young fish to avoid reef-based parasites.     

Brachiopods from cryptic coral reef habitats in the northern Red Sea

In contrast to the Palaeozoic to Jurassic fossil record, modern tropical and subtropical shallow-water brachiopods are typically small-sized and mostly restricted to cryptic habitats in coral reefs, but information on microhabitat-composition is scant. At Dahab, northern Red Sea, living brachiopods of the genus Argyrotheca were only detected on massively encrusted coral colonies attached to encrusting foraminifers and coralline red algae. Three samples from autochthonous sediments underneath coral colonies are comparatively rich in the brachiopod genera Megerlia and Argyrotheca, and additionally show low numbers of Novocrania and Thecidellina. Based on a coarse-grain analysis including more than 16,000 components >1 mm, these brachiopod shells co-occur with skeletal components of 11 higher taxa. Decapods, fixosessile foraminifers, molluscs, scleractinians, and coralline red algae clearly dominate the assemblages. Brachiopods in this study always contribute less than 2% to the sediment composition. This confirms previous results that even in brachiopod habitats the contribution of brachiopod shells to the total sediment composition is almost negligible. Our study indicates that brachiopods co-occur with pteriomorph bivalves and other epifauna in the cryptic habitats with limited space for encrusters or epibionts on the undersides of scleractinians and it tentatively supports the hypothesis of brachiopods preferring habitats with low grazing pressure, because shelly components of grazers (polyplacophorans and regular echinoids) are rare in our samples.