Haliotis Midae Research Articles

Domain repeats related to innate immunity in the South African abalone, Haliotis midae

Molluscs predominately use the cellular defence system as the primary mechanism of defence against pathogenic infection, in which haemocytes play a pivotal role. Haliotis midae is a commercially important South African species that it is susceptible to bacterial pathogens, fungal and yeast infections in the farming environment. The current study aims to enrich the current knowledge regarding H. midae innate immunity by investigating the presence and evolution of domain repeats. The bioinformatics approach used in this study, detected five repeat families in the H. midae transcriptome. These repeats families include mixed alpha and beta (leucine-rich and ankyrin), spectrin repeats, beta-propellers (WD40) and alfa-structure repeat (TPR-like). The expansion of key gene families related to host defence may be important to abalone adaptation to life in a pathogen-rich environment.

Transcriptome profiles of wild and cultured South African abalone, Haliotis midae

This report describes the use of pyrosequencing technologies to generate the first comparative analysis of de novo assembled transcriptome data from cultured and wild specimens of the South African abalone. The transcriptome data and database described here provide a significant genomic resource for abalone research. The data set annotated 11,240 genes, which matched genes with known functions in other species. A large number of transmembrane protein domains (4087) that may indicate a high portion of undiscovered gene receptors were identified. Further, we detected an interesting set of transcription factors (516) that are valuable candidates for participating in regulatory events in developmental (such as cell proliferation and differentiation) and reproductive processes.

A chain of fools: or, why it is so hard to stop overfishing

We fish too much, and by doing so, we threaten marine ecosystems and people’s livelihoods. But the curious thing is: we have known this for a long time. Nonetheless, we continue to overfish. How is that possible? Why can we not stop? This paper recounts our search for an answer. We start by giving an overview of how scientists explain overfishing, and suggest that the riddle of its obduracy has not been addressed systematically. We conceptualize overfishing as an unplanned and unintended outcome of a chain of interrelated social and ecological events. We then analyze the chain of events leading to overfishing for two typical cases – the groundfish fishery in the Gulf of Maine and the South African abalone (Haliotis midae, Haliotidae) fishery – and two atypical cases where overfishing has stopped or been substantially reduced – the Patagonian and Antarctic toothfish (Dissostichus eleginoides, Nototheniidae and Dissostichus mawsoni, Nototheniidae) fishery in the Southern Ocean, and the Atlantic cod (Gadus morhua, Gadidae) fishery in the Barents Sea. Studying and comparing these cases reveals no sufficient set of factors to explain the persistence of overfishing. Rather, distinct pathways emerge from a concatenation of proximate and remote factors, leading to and sustaining overfishing. Understanding these pathways and their mechanisms can assist in locating leverage points for intervention aimed to stop overfishing.

Polydorid polychaetes (Spionidae) on farmed and wild abalone (Haliotis midae) in South Africa: an epidemiological survey

Although there has been an increase in our understanding of the shell-boring polydorids that infest abalone Haliotis midae in South Africa, abalone from a limited number of farms, and wild populations from east of Cape Agulhas only, have been examined. To gain further knowledge and a more complete understanding of the local distribution of polydorids, we examined up to 30 abalone from each of 14 farms in the Northern, Western and Eastern Cape provinces, and five wild sites in the Western Cape, west of Cape Agulhas. Farm and wild communities were significantly different from each other (ANOSIM, r = 0.632, p < 0.002) and Bray–Curtis cluster analysis showed that most farms clustered separately from wild sites. Boccardia proboscidea was present on all but one of the infested farms, whereas some were also infested by Dipolydora capensis. By contrast, D. capensis was present at all wild sites sampled, whereas B. proboscidea was absent from these sites. Polydora hoplura, a common shell-boring pest, was present at only two farms and two wild sites. There appeared to be some exchange of polydorids among farms and between farms and wild abalone. The farm on the East Coast did not cluster with any other sites, indicating a unique composition of polydorids on that coast. No new, potentially problematic, species was encountered, although four species showed an increase in their known distribution.

Rainbow trout silage oil as immunity enhancing feed ingredient in formulated diets for South African abalone Haliotis midae

Abstract This study was conducted to evaluate the effects of rainbow trout silage oil on the immune function, growth performance and foot muscle fatty acid profile and proximate composition of South African abalone Haliotis midae . Rainbow trout silage oil was prepared from farmed rainbow trout processing waste, included at 25 g kg − 1 in a formulated diet and compared to a control diet in a 153 day growth trial. It was found that rainbow trout silage oil addition significantly improved abalone cellular immune function by increasing hemocyte phagocytic activity from 48.8% in the control diet to 65.2% in the silage oil diet. However, production performance was negatively impacted: specific growth rate (SGR) and final animal weight were decreased 17% and 5.6% respectively, and feed conversion ratio (FCR) increased by 25.1%. Foot muscle fatty acid profile and proximate composition were unaffected. It is concluded that dietary fish silage oil inclusion can improve cellular immune function in H. midae , but that the optimal inclusion level should be determined in order to negate the negative effects on production efficiency.

The effects of protein hydrolysates on the immunity and growth of the abalone Haliotis midae

Abstract Dietary hydrolysed proteins have been shown to stimulate the non-specific immunity of various finfish species, whilst the potential for immune stimulation of these feed ingredients in abalone has not been investigated. The immune-stimulating potential of two hydrolysed protein sources (self-prepared fish silage and a commercial fish protein hydrolysate) at two dietary inclusion levels in abalone diets was therefore measured in the South African abalone Haliotis midae , using animals of initial mean weight of 123 g–128 g. Diets containing the high inclusion levels were also fed in two feeding regimes: continuous feeding, or phase feeding, where the hydrolysed protein diets were alternated monthly with the control diet. In low inclusion diets, hydrolysate inclusion contributed 6 g·kg − 1 protein in the final diet, whilst in high inclusion diets this was increased to 18 g·kg − 1 . It was found that the low inclusion level of the commercial hydrolysate significantly increased the cellular immunity through increasing the phagocytic activity of haemocytes by 18% compared to the control diet, whilst none of the other diets showed any significant differences compared to the control. Both inclusion levels of the commercial hydrolysed protein significantly improved daily weight increase of animals. Phase feeding had no positive impacts on immunity or production performance. The inclusion of both types of hydrolysed protein sources significantly decreased the water stability of feeds compared to the control, except at the lowest inclusion level of the commercial hydrolysate. It is concluded that the use of dietary hydrolysed proteins can lead to improved cellular immunity and growth in abalone, however it is important to determine appropriate inclusion levels to prevent negative impacts on feed water stability and production performance.

Comparison of population genetic estimates amongst wild, F1 and F2 cultured abalone (Haliotis midae)

Haliotis midae is South Africa's most important aquaculture species. The reproduction cycle is currently not closed as many farms rely on wild-caught broodstock for seed production. However, there is an increasing interest in genetic improvement in commercial stocks, with a growing number of producers implementing selective breeding strategies. High throughput commercial production and mass spawning make it difficult to maintain breeding records; therefore, mostly mass selection is practised. The high fecundity and unequal parental contributions also often lead to increased levels of inbreeding. This study therefore aimed to assess the genetic effects of such breeding practices on commercial populations of H. midae. Using microsatellite loci, the genetic properties of a wild, an F1 and an F2 population were estimated and compared. Although there was no significant loss of genetic diversity amongst the cultured populations in comparison with the wild progenitor population, there was low-to-moderate genetic differentiation between populations. Relatedness amongst the F2 population was significant, and the rate of inbreeding was high. The effective population size for the F2 (±50) was also comparatively small with respect to the wild (∞) and F1 (±470) populations. These results suggest that farms need to give caution to breeding practices beyond the first (F1) generation and aim to increase effective population sizes and minimise inbreeding to ensure long-term genetic gain and productivity. This study also confirms the usefulness of population genetic analyses for commercial breeding and stock management in the absence of extensive pedigree records.

Detection and Localisation of the Abalone Probiotic Vibrio midae SY9 and Its Extracellular Protease, VmproA, within the Digestive Tract of the South African Abalone, Haliotis midae

Probiotics have been widely reported to increase the growth rate of commercially important fish and shellfish by enhancing the digestion of ingested feed through the production of extracellular enzymes such as proteases and alginases. In order to investigate this further, the objective of this study was to localise the bacterial probiont Vibrio midae SY9 and one of the extracellular proteases it produces in the digestive tract of the South African abalone Haliotis midae. This was accomplished by inserting a promotorless gfp gene into the chromosome of the bacterium which was incorporated in an artificial, fishmeal-based abalone feed. In situ histological comparison of abalone fed either a basal diet or the basal diet supplemented with V. midae SY9::Tn10.52 using a cocktail of DNA probes to the gfp gene localised the probiont to the crop/stomach and intestinal regions of the H. midae digestive tract. Generally, the ingested probiotic bacterium occurred in association with feed and particulate matter within the crop/stomach and intestinal regions, as well as adhered to the wall of the crop/stomach. Histological immunohistochemical examination using polyclonal anti-VmproA antibodies localised an extracellular protease produced by V. midae SY9 to the H. midae crop/stomach and intestine where it appeared to be associated with feed and/or other particulate matter in the abalone gut. Thus the data suggests that V. midae SY9 colonises and/or adheres to the mucous lining of the abalone gut. Furthermore, the close association observed between the bacterium, its extracellular protease and ingested feed particles supports the theory that V. midae SY9 elevates in situ digestive enzyme levels and thus enhances feed digestion in farmed abalone.

Was overfishing of predatory fish responsible for a lobster-induced regime shift in the Benguela?

The top-down effects of predators have been demonstrated for terrestrial, freshwater and marine systems and their removal can cause a shift in ecosystem state. In many cases, the depletion of top-predators occurred long before humans began monitoring these systems, but models can elucidate likely ecosystem changes. In this paper we use a multispecies model to demonstrate that the abundance of predators can induce different ecosystem states, emphasising the importance of sustainable harvesting. Our model is founded on empirical data documenting a regime-shift that took place in the kelp-forest ecosystem along the south-west coast of South Africa following an ‘invasion’ by the West Coast rock lobster Jasus lalandii in the early 1990s. Formerly dominated by herbivores and encrusting corallines, the ecosystem became dominated by lobsters and macroalgae. A combination of illegal fishing of abalone Haliotis midae and the lobster-induced extermination of the urchin Parechinus angulosus (which shelters juvenile abalone) depleted abalone stocks severely. In this currently over-fished ecosystem where predators of lobsters are scarce, the regime-shift appears irreversible. Using an intermediate complexity model of lobster, urchin and abalone relationships, we simulated the effects of predatory fish at various population levels on the course of development of the currently lobster-dominated system. Our results indicate that current levels of fish biomass (<10% of the pristine population) would have been too small to significantly affect the lobster population. However, at pristine levels of fish abundance (or even 50% less), the invasion of lobsters would have been controlled by linefish, and urchin and abalone populations would not have collapsed. The effect of the lobster invasion had a greater influence on abalone abundance than illegal fishing, but together they severely depleted the abalone. Ecosystem effects of historical exploitation of top predators are of central concern, and our ecosystem model indicates that depletion of linefish had severe consequences for lobsters, urchins and abalone, illustrating how overfishing of top predators may precipitate regime shifts, with profound socio-economic consequences.

A decade of illegal fishing in Table Mountain National Park (2000–2009): trends in the illicit harvest of abalone Haliotis midae and West Coast rock lobster Jasus lalandii

Illegal fishing activities are reported to be on the increase in South Africa, including in its marine protected areas (MPAs). Research is presented on the nature and the scale of illegal fishing in Table Mountain National Park (TMNP) by analysing the numbers of abalone Haliotis midae and West Coast rock lobster Jasus lalandii confiscated from fishers operating in the park's marine protected area between 2000 and 2009. Data were collected from offence logbooks maintained by South African National Parks rangers and managers, the South African Police Services, and interviews with alleged or self-confessed illegal fishers. The research findings indicate that the annual numbers of illegally fished abalone and rock lobsters have increased significantly over time. Spatial analysis suggests that confiscations of abalone occur predominantly on the east coast of the park, whereas higher confiscations of illegally fished rock lobsters occur on the west coast. It is clear from this research that new and more efficient approaches will need to be designed and implemented to minimise illegal fishing in the TMNP MPA. Context-specific conservation targets that acknowledge and integrate social as well as developmental needs are required, and may be essential for limiting biodiversity loss in the longer term, which will ultimately ensure the success of fisheries management and conservation in TMNP.

An experimental test of the effect of the abalone Haliotis midae on benthic community composition

The South African abalone Haliotis midae is a commercially valuable species, but its numbers are seriously depleted due to illegal fishing. Overfishing not only affects the targeted species, but also potentially influences the integrity and functioning of the ecosystem through associated changes in community composition. We assessed the herbivorous effects of H. midae on the benthos through a 6-month field experiment involving exclusion/inclusion plots. This showed that benthic community composition remained largely unchanged whether H. midae was present or absent and specifically revealed no evidence that H. midae diminishes algal biomass, probably because in the region where we worked, it feeds mainly by trapping drift kelp rather than grazing on attached algae. The absence of responses to the removal of abalone was additionally evident in comparison with indices of abundance and diversity, functional groups and individual species. In the absence of abalone, there were subtle increases in the green alga Codium stephensiae and the crustose brown alga Zeacarpa leiomorpha and decreases in the four species of foliose macroalgae, but none of these effects were significant. The presence of H. midae thus had no negative grazing effects on any elements of the epilithic community composition. It does, however, have other influences, including occupation of space and provision of habitat in the form of its shell, which supports a community significantly different from the surrounding rocks.

Oxidative stress in abalone: The role of temperature, oxygen and l-proline supplementation

In their natural distribution range and in on land-based mariculture facilities, Haliotis midae may be exposed to a relatively wide range of temperatures and oxygen levels, that can induce differential degrees of stress with concomitant metabolic responses.In the current study juvenile H. midae were exposed for one month to two constant temperatures (14°C and 19°C), with three oxygen levels (82%, 98% and 126% oxygen saturation) at each temperature. One additional treatment was included at 19°C, 126% oxygen, where feed was supplemented with l-proline. The results presented here suggest that long-term 19°C exposure is more stressful to H. midae juveniles than a 14°C exposure of the same duration, since exposure to higher temperature generally resulted in higher activities of the antioxidant enzymes measured but lower total antioxidant capacities. Haemolymph glucose levels responded to oxygen treatment at the 14°C exposure, but the same response was not initiated in the 19°C treatment, possibly because the animals exposed to the 19°C treatment relied on proteins to a larger extent.Increased oxygen levels at the 19°C treatment did not cause more DNA damage or protein carbonylation compared to lower oxygen levels. Supplementing feed with l-proline at the high temperature/high oxygen treatment caused a decrease in haemolymph haemocyanin (which may exhibit catalase function) content and a slight decrease in some of the antioxidant enzyme activities, while maintaining the same total antioxidant capacity compared with the same treatment where feed was not supplemented. The overall DNA integrity profile also changed with proline-supplementation, as did the induction level of hsp70. When projecting tissue growth over a 12month period under these conditions, we estimate that H. midae juveniles in this study may exhibit a significant increase in mass when their feed was supplemented with l-proline, indicating that l-proline supplementation provides some benefit to the individual. It might therefore warrant conducting long-term growth trials on land-based mariculture facilities to further investigate its potential benefit to the mariculture industry.

Transcriptome-Wide Single Nucleotide Polymorphisms (SNPs) for Abalone (Haliotis midae): Validation and Application Using GoldenGate Medium-Throughput Genotyping Assays

Haliotis midae is one of the most valuable commercial abalone species in the world, but is highly vulnerable, due to exploitation, habitat destruction and predation. In order to preserve wild and cultured stocks, genetic management and improvement of the species has become crucial. Fundamental to this is the availability and employment of molecular markers, such as microsatellites and single nucleotide (SNPs). Transcriptome sequences generated through sequencing-by-synthesis technology were utilized for the in vitro and in silico identification of 505 putative SNPs from a total of 316 selected contigs. A subset of 234 SNPs were further validated and characterized in wild and cultured abalone using two Illumina GoldenGate genotyping assays. Combined with VeraCode technology, this genotyping platform yielded a 65%–69% conversion rate (percentage polymorphic markers) with a global genotyping success rate of 76%–85% and provided a viable means for validating SNP markers in a non-model species. The utility of 31 of the validated SNPs in population structure analysis was confirmed, while a large number of SNPs (174) were shown to be informative and are, thus, good candidates for linkage map construction. The non-synonymous SNPs (50) located in coding regions of genes that showed similarities with known proteins will also be useful for genetic applications, such as the marker-assisted selection of genes of relevance to abalone aquaculture.

Metabolic readjustment in juvenile South African abalone (Haliotis midae) acclimated to combinations of temperature and dissolved oxygen levels

When ectotherms are exposed to high temperatures, cellular oxygen demand increases. When oxygen demand exceeds oxygen availability, animals may experience functional hypoxia regardless of environmental oxygen levels.In the current study, we aimed to assess if oxidative or thermal restrictions exist when winter-acclimated Haliotis midae juveniles are re-acclimated to temperatures within their thermal tolerance range, but with one temperature outside the average daily temperatures they would experience in winter. At each acclimation temperature (14, 16 and 19°C), we also manipulated dissolved oxygen levels to obtain oxygen saturation levels of 82%, 98% and 126%.Our results indicate growth restrictions and hypometabolism at all oxygen treatments at 19°C. Efficiency of mitochondrial respiration, as indicated by an increase in cytochrome c oxidase subunits 1 and 2 (cox1 and cox2) transcript level, was improved at 19°C, 98% oxygen. A change in substrate utilisation, from a glycogen/lipid-driven metabolism to almost exclusively protein-driven metabolism provided the fuel for more effective cellular respiration. A further decrease in metabolic rate at 19°C, 126% oxygen, this time without an increase in cox1 and cox2 transcripts, may indicate some improvement in gas exchange, but without improvement in growth.We propose that 19°C is nearing a long-term limiting temperature for winter-acclimated SA abalone, and that animals survive this condition by re-allocating available energy to survival mechanisms instead of growth. This species, already threatened by over-harvesting, may be further threatened by increases in surface seawater temperatures.

Development of an experimental laboratory fertilization protocol for the South African abalone, Haliotis midae (Linnaeus 1758)

In this study, effective gamete concentrations, egg viability, and fertilization volumes were evaluated for Haliotis midae (L.). Sperm concentrations between 5 × 103 and 5 × 104 mL−1 (p > 0.05) consistently resulted in high hatch-out rates (96 ± 1%). At concentrations higher than 5 × 105 mL−1, hatch-out rates decreased to 69 ± 7% (p < 0.05). Egg concentrations between 10 and 125 eggs mL−1 resulted in high fertilization rates, with 50 eggs mL−1 being the ideal concentration for fertilization in H. midae. Egg viability was consistently high up to 100 min post-spawning, with a decrease in hatch-out success, when eggs were fertilized 120 min post-spawning. Fertilization volumes did not affect successful hatch-out. The results from this study can be implemented by South African abalone farms to increase hatch-out rates and subsequent culture. It can also be used as basis for the development of fertilization protocols in other marine invertebrate species.

Detection of molecular signatures of selection at microsatellite loci in the South African abalone (Haliotis midae) using a population genomic approach

Identifying genomic regions that may be under selection is important for elucidating the genetic architecture of complex phenotypes underlying adaptation to heterogeneous environments. A population genomic approach, using a classical neutrality test and various Fst-outlier detection methods was employed to evaluate genome-wide polymorphism data in order to identify loci that may be candidates for selection amongst six populations (three cultured and three wild) of the South African abalone, Haliotis midae. Approximately 9% of the genome-wide microsatellite markers were putatively subject to directional selection, whilst 6–18% of the genome is thought to be influenced by balancing selection. Genetic diversity estimates for candidate loci under directional selection was significantly reduced in comparison to candidate neutral loci, whilst candidate balancing selection loci demonstrated significantly higher levels of genetic diversity (Kruskal–Wallis test, P<0.05). Pairwise Fst estimates based on candidate directional selection loci also demonstrated increased levels of differentiation between study populations. Various candidate loci under selection showed significant inter-chromosomal linkage disequilibrium, suggesting possible gene-networks underling adaptive phenotypes. Furthermore, several loci had significant hits to known genes when performing BLAST searches to NCBI's non-redundant databases, whilst others are known to be derived from expressed sequences even though homology to a known gene could not be established. A number of loci also demonstrated relatively high similarity to transposable elements. The association of these loci to functional and genomically active sequences could in part explain the observed signatures of selection.

Integrated Linkage Map ofHaliotis midaeLinnaeus Based on Microsatellite and SNP Markers

The South African abalone Haliotis midae Linnaeus is the most important aquaculture species in South Africa. Marker-assisted selection is envisioned to play an integral part of the genetic improvement program of abalone, and therefore the generation of linkage maps for quantitative trait loci analyses are necessary. This study reports on a first-generation linkage map for H. midae based on microsatellite and single nucleotide polymorphism (SNP) markers. Ten full-sib families were screened with a total of 508 molecular markers derived from genomic and expressed sequence tag sequences. Linkage maps were constructed for each of the families and combined to create an integrated linkage map. The integrated linkage map consists of 186 markers that included 116 microsatellites and 70 SNPs. These markers mapped to 18 linkage groups, which corresponds to the haploid chromosome number of H. midae. The average genome length was estimated at 1,312 cM, displaying an average marker interval of 6.88 cM with 80% genome coverage. The female map is 1.16-fold longer than the male map, indicating differences in recombination rate between the sexes. The association of markers with known genes as well as with transposon elements was also investigated. This study resulted in the first linkage map constructed for any haliotid in which both microsatellite and SNP markers were used, and the results provide a framework for future applications in quantitative trait loci identification.

Reproductive Histology of CulturedHaliotis midae(Linnaeus, 1758) and Preliminary Evaluation of Maturation

This study reports the first classification of cytological structures in developing gonads of cultivated Haliotis midae. Gonad tissue from animals ranging from 6–48 mo of age was evaluated histologically. The process of oogenesis was divided into 9 distinct stages according to the size and histological structures of each cell type, whereas spermatogenesis was divided into 5 distinct stages. Six-mo-old animals showed initial development of germinal epithelium, whereas at 12 mo of age, the gonad cavity was visible. In 18-mo-old males, spermatogonia and primary spermatocytes were identified, whereas in 24-mo-old males, all cell stages of spermatogenesis were present. Oogonia and stage 1 oocytes were seen in the gonad cavity of an 18-mo-old female, whereas in the gonad of 2-y-old females, all stages of oogenesis could be distinguished, although mature oocytes were not abundant.

Modeling a Regime Shift in a Kelp Forest Ecosystem Caused by a Lobster Range Expansion

The South African West Coast rock lobster, Jasus lalandii (H. Milne-Edwards, 1837), has expanded its range to the southeast, where its abundance has increased radically. The ecological consequences of this are likely to be considerable. We employed a minimally realistic model to simulate the and to explore interactions of J. lalandii with the sea urchin, Parechinus angulosus (Leske, 1778), and the abalone, Haliotis midae Linnaeus, 1758, juveniles of which shelter beneath this urchin. Model fits to empirical data were good, although species-interaction terms were difficult to estimate. Base-case trajectories indicated: (1) Lobster biomass peaked at about 1000 t in 1994 and was then reduced by fishing to a stable value approximately 50% lower by 2008. (2) Urchins remained close to carrying capacity in areas but collapsed to local extinction by 1997 in the area. (3) Abalone declined over 2000-2008 in noninvaded areas because of illegal fishing and collapsed to near zero in the invaded area because of illegal fishing combined with increased lobster abundance. Sensitivity analyses favored the hypothesis that the invasion was due to adult immigration rather than larval recruitment. Modeled 50-yr projections indicated that urchins will remain locally extinct in the area, even 50 yrs into the future. The abalone collapse in the area would persist >50 yrs, even if lobsters were absent. We argue that the lobster triggered an alternative stable state, making a return to pre-invasion conditions unlikely.

Comparing community structure on shells of the abalone Haliotis midae and adjacent rock: implications for biodiversity

This paper concerns the effects on biodiversity of depletion of the South African abalone Haliotis midae, which is a long-lived species with a large corrugated shell that provides a habitat for diverse benthic organisms. We compared community structure on H. midae shells with that on adjacent rock at three sites (Cape Point and Danger Point sites A and B) and at two different times of the year at one of these sites. Shells of H. midae consistently supported communities that were distinctly different from those on rock. In particular, three species of non-geniculate (encrusting) corallines, Titanoderma polycephalum, Mesophyllum engelhartii and Spongites discoideus, were all found either exclusively or predominantly on shells, whereas another non-geniculate coralline, Heydrichia woelkerlingii, occurred almost exclusively on adjacent rock. The primary rocky substratum, however, supported a higher number of species than abalone shells. Possible reasons for the differences between the two substrata include the relative age, microtopography and hardness of the substrata; the abundance of grazers on them; and the relative age of different zones of the abalone shell, which support communities at different stages of succession. Diversity on shells was lowest in zones that were either very young or very old, in keeping with the intermediate disturbance hypothesis. The distinctiveness of shell epibiota will increase β diversity despite having a lower α diversity than that of adjacent rock. Decimation of H. midae by overfishing therefore has implications for biodiversity conservation.