Abundance and clonal replication in the tropical corallimorpharian Rhodactis rhodostoma

Abstract. The corallimorpharian Rhodactis rhodostoma appears to be an opportunistic species capable of rapidly monopolizing patches of unoccupied shallow substrate on tropical reefs. On a fringing coral reef at Eilat, Israel, northern Red Sea, we examined patterns of abundance and clonal replication in R. rhodostoma in order to understand the modes and rates of spread of polyps across the reef flat. Polyps were abundant on the inner reef flat (maximum 1510 polyps m⁻² and 69% cover), rare on the outer reef flat, and completely absent on the outer reef slope at >3 m depth. Individuals cloned throughout the year via 3 distinct modes: longitudinal fission, inverse budding, and marginal budding. Marginal budding is a replicative mode not previously described. Cloning mode varied significantly with polyp size. Approximately 9% of polyps cloned each month, leading to a clonal doubling time of about 1 year. The rate of cloning varied seasonally and depended on day length and seawater temperature, except for a brief reduction in cloning during midsummer when polyps spawned gametes. Polyps of R. rhodostoma appear to have replicated extensively following a catastrophic low-tide disturbance in 1970, and have become an alternate dominant to stony corals on parts of the reef flat.     

Size-structural shifts reveal intensity of exploitation in coral reef fisheries

Fisheries exploitation represents a considerable threat to coral reef fish resources because even modest levels of extraction can alter ecological dynamics via shifts of stock size, species composition, and size-structure of the fish assemblage. Although species occupying higher trophic groups are known to suffer the majority of exploitative effects, changes in composition among lower trophic groups may be major, though are not frequently explored. Using size-based biomass spectrum analysis, we investigate the effects of fishing on the size-structure of coral reef fish assemblages spanning four geopolitical regions and determine if patterns of exploitation vary across trophic groups. Our analyses reveal striking evidence for the variety of effects fisheries exploitation can have on coral reef fish assemblages. When examining biomass spectra across the entire fish assemblage we found consistent evidence of size-specific exploitation, in which large-bodied individuals experience disproportionate reductions. The pattern was paralleled by and likely driven by, strongly size-specific reductions among top predators. In contrast, evidence of exploitation patterns was variable among lower trophic groups, in many cases including evidence of reductions across all size classes. The breadth of size classes and trophic groups that showed evidence of exploitation related positively to local human population density and diversity of fishing methods employed. Our findings highlight the complexity of coral reef fisheries and that the effects of exploitation on coral reefs can be realized throughout the entire fish assemblage, across multiple trophic groups and not solely restricted to large-bodied top-predators. Size-specific changes among fishes of lower trophic groups likely lead to altered ecological functioning of heavily exploited coral reefs. Together these findings reinforce the value of taking a multi-trophic group approach to monitoring and managing coral reef fisheries.     

Factors influencing algal species assemblages on reef and cobble substrata off Ghana

Subtidal algal assemblages were studied on two substrata, rocky reefs and calcareous cobbles. Although the two occur together at comparable depths, their vegetation differs in species composition and richness, and in patterns of plant size, life form, and longevity. The reef bears a species-rich, patchy cover of small filamentous and crustose forms, with occasional clumps of more robust species. The cobbles support a sparse cover of large leafy and dendritic species in addition to many of the smaller species found on the reef. The floristic separation arises from differential establishment and survival of species under conditions of (1) grazing by fish and urchins (on the reef only), and (2) seasonal physical disturbance during storms leading to the removal of most algae (on the cobbles only). Both substrata show a seasonal floristic cycle, but the trend is more pronounced on cobbles. Species do not depart from randomness in their patterns of co-occurrence on individual cobbles or reef fragments. Interspecific competition appears comparatively unimportant in determining species composition on either substratum.     

Fishing impacts on the marine inorganic carbon cycle

1.  Teleost fish excrete precipitated carbonate and make significant contributions to the marine inorganic carbon cycle at regional and global scales. As total carbonate production is linked to fish size and abundance, fishing is predicted to affect carbonate production by modifying fish abundance and size-structure.
2.  We draw on concepts from physiology, metabolic ecology, life history theory, population dynamics and community ecology to develop, validate and apply analytical tools to assess fishing impacts on carbonate production. Outputs suggest that population and community carbonate production fall rapidly at lower rates of fishing than those used as management targets for sustainable yield.
3.  Theoretical predictions are corroborated by estimated trends in carbonate production by a herring population and a coral reef fish community subject to fishing. Our analytical results build on widely applicable relationships between life history parameters and metabolic rates, and can be generalized to most fished ecosystems.
4.   Synthesis and applications . If the maintenance of chemical processes as well as biological process were adopted as a management objective for fisheries then the methods we have developed can be applied to assess the effects of fishing on carbonate production and to advise on acceptable rates of fishing. Maintenance of this ecosystem service would require lower rates of fishing mortality than those recommended to achieve sustainable yield.     

Effects of variation in resource acquisition during different stages of the life cycle on life‐history traits and trade‐offs in a burying beetle

Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.     

Sexual reproduction in the tropical corallimorpharian Rhodactis rhodostoma

Abstract. Polyps of the tropical corallimorpharian Rhodactis rhodostoma segregate sexes between center and edge positions within aggregations produced by clonal replication. On a reef flat at Eilat, northern Red Sea, infertile polyps and males occur mainly along the edges of clonal aggregations, while females mostly occupy central positions within each aggregation. In addition, on the inner to middle reef flat where polyps of this species are abundant, aggregations consist mostly of females. On the outer reef flat, where polyps are rare, a sampled aggregation consisted mostly of males and infertile polyps. Male polyps are significantly smaller than females, and the smallest polyps are infertile. Fecundity increases significantly with polyp size in females, but testis size and number do not vary with body size in males. Oocytes are present in polyps during most of the year and gradually increase in size until annual spawning in June-July during the period of maximum day length. Testes do not vary significantly in size during the year and remain a small proportion of body mass (>8%). In contrast, females invest up to 30% of their body mass into gonads during the months immediately before spawning. The annual spawning of gametes coincides with a temporary drop in the frequency of clonal replication by polyps. We estimate that each female polyp of R. rhodostoma may release up to 3000 large eggs (500 μm in maximum diameter) each summer. The high investment of this corallimorpharian in sexual production of planktonic propagules may allow rapid dispersal to reef habitats distant from parent populations.     

Mechanical vulnerability explains size‐dependent mortality of reef corals

Understanding life history and demographic variation among species within communities is a central ecological goal. Mortality schedules are especially important in ecosystems where disturbance plays a major role in structuring communities, such as coral reefs. Here, we test whether a trait‐based, mechanistic model of mechanical vulnerability in corals can explain mortality schedules. Specifically, we ask whether species that become increasingly vulnerable to hydrodynamic dislodgment as they grow have bathtub‐shaped mortality curves, whereas species that remain mechanically stable have decreasing mortality rates with size, as predicted by classical life history theory for reef corals. We find that size‐dependent mortality is highly consistent between species with the same growth form and that the shape of size‐dependent mortality for each growth form can be explained by mechanical vulnerability. Our findings highlight the feasibility of predicting assemblage‐scale mortality patterns on coral reefs with trait‐based approaches.     

Growth and population dynamic model of the reef coral Fungia granulosa Klunzinger, 1879 at Eilat, northern Red Sea

The lack of population dynamic information for most species of stony corals is due in part to their complicated life histories that may include fission, fusion and partial mortality of colonies, leading to an uncoupling of coral age and size. However, some reef-building corals may produce compact upright or free-living individuals in which the above processes rarely occur, or are clearly detectable. In some of these corals, individual age may be determined from size, and standard growth and population dynamic models may be applied to gain an accurate picture of their life history. We measured long-term growth rates (up to 2.5 years) of individuals of the free-living mushroom coral Fungia granulosa Klunzinger, 1879 at Eilat, northern Red Sea, and determined the size structure of a population on the shallow reef slope. We then applied growth and population models to the data to obtain estimates of coral age, mortality rate, and life expectancy in members of this species. In the field, few F. granulosa polyps suffered partial mortality of >10% of their tissues. Thus, the majority of polyps grew isometrically and determinately, virtually ceasing growth by about 30-40 years of age. Coral ages as revealed by skeletal growth rings were similar to those estimated from a growth curve based on field data. The frequency of individuals in each age class on the reef slope decreased exponentially with coral age, indicating high mortality rates when corals were young. The maximum coral age observed in the field population (31 years) was similar to that estimated by application of a population dynamic model (30 years). Calculated rates of growth, mortality and life expectancy for F. granulosa were within the range of those known for other stony corals. Our results reveal a young, dynamic population of this species on Eilat reefs, with high turnover rates and short lifespans. Such information is important for understanding recovery of coral reefs from disturbances, and for application to the management of commercially exploited coral populations.     

Implications of body size for interspecific interactions and assemblage organization among coral-reef fishes

Abstract Population size-structure is often ignored in assemblage-level studies of reef fishes, which usually rely on static and dynamic patterns of relative total abundance to infer what mechanisms organize those assemblages. However, body size has substantial effects on processes that affect competitive relationships between species: (i) small, recently recruited fish, which usually(?) suffer high mortality, can dominate total abundance and strongly influence the dynamics of the relative total abundances of different species, while having little effect on interspecific biomass relations; (ii) numeric abundance and biomass of a species can vary independently, due to habitat variation in population size-structure resulting from variation in mortality and growth, as well as habitat selection; and (iii) population size-structure affects the potential for and outcome of interspecific competition due to (a) ontogenetic change in types of resources used, (b) levels of resource needs being dependent on individual and species biomass rather than numbers, (c) advantages due to large size in behavioural contests, (d) variation in population size-structure being linked to habitat preference, which affects expression of competitive dominance, and (e) size dependency in the development of interspecific resource-sharing relationships. Assemblage-level analyses that ignore such size effects may fail to detect important effects of interspecific interactions.     

Recurrent disturbances and the degradation of hard coral communities in taiwan

Recurrent disturbances can have a critical effect on the structure and function of coral reef communities. In this study, long-term changes were examined in the hard coral community at Wanlitung, in southern Taiwan, between 1985 and 2010. In this 26 year interval, the reef has experienced repeated disturbances that include six typhoons and two coral-bleaching events. The frequency of disturbance has meant that species susceptible to disturbance, such as those in the genus Acropora and Montipora have almost disappeared from the reef. Indeed, almost all hard coral species have declined in abundance, with the result that total hard coral cover in 2010 (17.7%) was less than half what it was in 1985 (47.5%). In addition, macro-algal cover has increased from 11.3% in 2003 to 28.5% in 2010. The frequency of disturbance combined with possible chronic influence of a growing human population mean that a diverse reef assemblage is unlikely to persist on this reef into the future.     

Responses of coral reef fishes to past climate changes are related to life‐history traits

Coral reefs and their associated fauna are largely impacted by ongoing climate change. Unravelling species responses to past climatic variations might provide clues on the consequence of ongoing changes. Here, we tested the relationship between changes in sea surface temperature and sea levels during the Quaternary and present‐day distributions of coral reef fish species. We investigated whether species‐specific responses are associated with life‐history traits. We collected a database of coral reef fish distribution together with life‐history traits for the Indo‐Pacific Ocean. We ran species distribution models (SDMs) on 3,725 tropical reef fish species using contemporary environmental factors together with a variable describing isolation from stable coral reef areas during the Quaternary. We quantified the variance explained independently by isolation from stable areas in the SDMs and related it to a set of species traits including body size and mobility. The variance purely explained by isolation from stable coral reef areas on the distribution of extant coral reef fish species largely varied across species. We observed a triangular relationship between the contribution of isolation from stable areas in the SDMs and body size. Species, whose distribution is more associated with historical changes, occurred predominantly in the Indo‐Australian archipelago, where the mean size of fish assemblages is the lowest. Our results suggest that the legacy of habitat changes of the Quaternary is still detectable in the extant distribution of many fish species, especially those with small body size and the most sedentary. Because they were the least able to colonize distant habitats in the past, fish species with smaller body size might have the most pronounced lags in tracking ongoing climate change.     

Grazing by a small fish affects the early stages of a post-settlement stony coral

Short-term experiments were used to isolate the detrimental effects of grazer disturbance on young corals, and determine the stage of development at which recruits are no longer susceptible to this disturbance. Artificial substrata containing an algal matrix and coral recruits of different life stages were exposed to grazing by epilithic algal matrix (EAM) feeding combtoothed blennies, Salarias fasciatus. Single polyp recruits were vulnerable to grazer disturbance, while multi-polyp recruits (ca. 6–8 polyps) survived with evidence of minor damage in the form of tissue and polyp loss. The result indicates that blennies, although small and possessing weak dentition, can negatively influence the survival of young coral recruits. The protruding structure of micro-nubbins, representing juvenile corals were not damaged, suggesting that coral achieving that size and form can escape such damage. Communicated by Ecology Editor Prof. Peter Mumby     

Near-future ocean acidification enhances the feeding rate and development of the herbivorous juveniles of the crown-of-thorns starfish, <Emphasis Type="Italic">Acanthaster planci</Emphasis>

Population outbreaks of the corallivorous crown-of-thorns starfish, Acanthaster planci, are a major contributor to the decline in coral reef across the Indo-Pacific. The success of A. planci and other reef species in a changing ocean will be influenced by juvenile performance because the naturally high mortality experienced at this sensitive life history stage maybe exacerbated by ocean warming and acidification. We investigated the effects of increased temperature and acidification on growth of newly metamorphosed juvenile A. planci and their feeding rates on crustose coralline algae (CCA) during the initial herbivorous phase of their life history. The juveniles were exposed to three temperature (26, 28, 30 °C) and three pH (NIST scale: 8.1, 7.8, 7.6) levels in a flow-through cross-factorial experiment. There were positive but independent effects of warming and acidification on juvenile growth and feeding. Early juveniles were highly tolerant to moderate increases in temperature (+2 °C above ambient) with the highest growth at 30 °C. Growth and feeding rates of A. planci on CCA were highest at pH 7.6. Thus, ocean warming and acidification may enhance the success of A. planci juveniles. In contrast to its coral prey, at this vulnerable developmental stage, A. planci appears to be highly resilient to future ocean change. Success of juveniles in a future ocean may have carry-over effects into the coral-eating life stage, increasing the threat to coral reef systems.     

Habitat utilization by coral reef fish: implications for specialists vs. generalists in a changing environment

1. The impact of environmental disturbance and habitat loss on associated species is expected to be dependent on a species' level of specialization. We examined habitat use and specialization of coral reef fish from the diverse and ecologically important family Pomacentridae, and determined which species are susceptible to declines in coral cover due to disturbance induced by crown-of-thorns seastar (COTS, Acanthaster planci L.). 2. A high proportion of pomacentrid species live in association with live coral as adults (40%) or juveniles (53%). Adults of many species had strong affiliations with branching corals, while juveniles favoured plating growth forms, reflecting the sizes of refuge provided by coral types. 3. Juveniles of species that associated with coral had narrower niche breadths than adult conspecifics, due to associations with specific coral types. The especially high coral association and narrower niche breadth of juveniles suggest that the presence of live coral is crucial for many species during early life history, and that disturbance-induced coral loss may have serious flow-on effects on adult abundance. 4. Microhabitat availability was a poor predictor of fish species abundance. Significant correlations between coverage of coral types and abundance of five adults and two juvenile species were detected; however, these relationships explained <35% and <10% of the variation in abundance of adult and juvenile species, respectively. 5. Niche breadth explained 74% of the variation in species' mean response to coral decline and it is clear that disturbance has a greater impact on resource specialists, suggesting that increasing frequency and intensity of coral loss will cause reef fish communities to become dominated by habitat generalists at the expense of coral-dwelling specialists.     

Contrasting patterns of density‐dependent selection at different life stages can create more than one fast–slow axis of life‐history variation

There has been much recent research interest in the existence of a major axis of life‐history variation along a fast–slow continuum within almost all major taxonomic groups. Eco‐evolutionary models of density‐dependent selection provide a general explanation for such observations of interspecific variation in the "pace of life." One issue, however, is that some large‐bodied long‐lived “slow” species (e.g., trees and large fish) often show an explosive “fast” type of reproduction with many small offspring, and species with “fast” adult life stages can have comparatively “slow” offspring life stages (e.g., mayflies). We attempt to explain such life‐history evolution using the same eco‐evolutionary modeling approach but with two life stages, separating adult reproductive strategies from offspring survival strategies. When the population dynamics in the two life stages are closely linked and affect each other, density‐dependent selection occurs in parallel on both reproduction and survival, producing the usual one‐dimensional fast–slow continuum (e.g., houseflies to blue whales). However, strong density dependence at either the adult reproduction or offspring survival life stage creates quasi‐independent population dynamics, allowing fast‐type reproduction alongside slow‐type survival (e.g., trees and large fish), or the perhaps rarer slow‐type reproduction alongside fast‐type survival (e.g., mayflies—short‐lived adults producing few long‐lived offspring). Therefore, most types of species life histories in nature can potentially be explained via the eco‐evolutionary consequences of density‐dependent selection given the possible separation of demographic effects at different life stages.     

Determinants of social organization in a coral reef fish,the blue tang, <Emphasis Type="Italic">Acanthurus coeruleus</Emphasis>

Synopsis We examined the abundance of blue tang surgeonfish, Acanthurus coeruleus, in each of three social modes (schooling, territorial, and wandering) in relation to size class, ecological variables, population density and time of day to discern potential determinants of social organization. We found individuals from all three social modes in all four fringing reef habitats (back reef, flat, crest and spur and groove zones) at our main site. Territorial tang density was highest in the flat zone, lowest in the spur and groove zone and negatively related to adult damselfish density. A higher proportion of tangs formed schools in reef zones with the highest densities of territorial damselfishes (spur and groove, crest). In the back reef and flat zones, where damselfish densities were very low, tangs rarely formed schools. Tangs in the wandering mode were most abundant in the back reef. The density of territorial tangs did not change with time of day, but non-territorial tangs tended to wander more in the morning and to form schools more at midday. Small yellow-phase juveniles were always territorial, and the incidence of territoriality decreased in the larger size classes, while schooling and wandering increased. Among similar fringing reefs, the incidence of territoriality increased with increasing population density. These data suggest that life history stage, damselfish density, and conspecific population density are important determinants of blue tang social organization.     

Compensating for delayed hatching across consecutive life‐history stages in an amphibian

Environmental conditions experienced early in the ontogeny can have a strong impact on individual fitness and performance later in life. Organisms may counteract the negative effects of poor developmental conditions by developing compensatory responses in growth and development. However, previous studies on compensatory responses have largely ignored the effects that poor embryonic conditions could have during the later life stages. In this study, we examined the effects of artificially delayed development in early life over two later life history transitions by investigating the compensatory growth of larval moor frogs Rana arvalis in response to temperature variation during embryonic development, and the associated costs during the larval ′and postmetamorphic stages. Low temperature during embryonic stage lead to delayed hatching at smaller size. The groups with delayed embryonic development showed strong compensatory growth during the larval stage, and reached similar metamorphic size than the controls in a shorter time. However, the most strongly delayed group was not able to fully catch up the total development time. These compensatory responses were found in the absence of photoperiod cues indicating that the delay in embryonic development was sufficient to initiate the compensatory response in larval growth and development. No apparent costs of compensatory growth were detected in terms of morphology or locomotor performance at the juvenile stage. We found that compensatory responses can be activated as early as at the embryonic stage and extend over several consecutive life history transitions, mitigating the effects of poor conditions experienced early in development. Potential short‐term costs in natural environments and the occurrence of long‐term costs, which prevent the generalisation of a faster larval life style, are discussed.     

Ontogenetic and spatial variation in size‐selective mortality of a marine fish

Although body size can affect individual fitness, ontogenetic and spatial variation in the ecology of an organism may determine the relative advantages of size and growth. During an 8‐year field study in the Bahamas, we examined selective mortality on size and growth throughout the entire reef‐associated life phase of a common coral‐reef fish, Stegastes partitus (the bicolour damselfish). On average, faster‐growing juveniles experienced greater mortality, though as adults, larger individuals had higher survival. Comparing patterns of selection observed at four separate populations revealed that greater population density was associated with stronger selection for larger adult size. Large adults may be favoured because they are superior competitors and less susceptible to gape‐limited predators. Laboratory experiments suggested that selective mortality of fast‐growing juveniles was likely because of risk‐prone foraging behaviour. These patterns suggest that variation in ecological interactions may lead to complex patterns of lifetime selection on body size.     

Identifying the ichthyoplankton of a coral reef using DNA barcodes

Marine fishes exhibit spectacular phenotypic changes during their ontogeny, and the identification of their early stages is challenging due to the paucity of diagnostic morphological characters at the species level. Meanwhile, the importance of early life stages in dispersal and connectivity has recently experienced an increasing interest in conservation programmes for coral reef fishes. This study aims at assessing the effectiveness of DNA barcoding for the automated identification of coral reef fish larvae through large‐scale ecosystemic sampling. Fish larvae were mainly collected using bongo nets and light traps around Moorea between September 2008 and August 2010 in 10 sites distributed in open waters. Fish larvae ranged from 2 to 100 mm of total length, with the most abundant individuals being <5 mm. Among the 505 individuals DNA barcoded, 373 larvae (i.e. 75%) were identified to the species level. A total of 106 species were detected, among which 11 corresponded to pelagic and bathypelagic species, while 95 corresponded to species observed at the adult stage on neighbouring reefs. This study highlights the benefits and pitfalls of using standardized molecular systems for species identification and illustrates the new possibilities enabled by DNA barcoding for future work on coral reef fish larval ecology.     

Disturbance and the structure of coral reef fish communities on the reef slope

Three levels of physical disturbance were applied to corals in permanent 10x10 m quadrats along a section of fringing reef at Lizard Island on the Great Barrier Reef to investigate the response of fish assemblages. Tabular and corymbose corals were overturned and left in situ, reducing total hard coral cover from ˜55% to ˜47%, ˜43%, and ˜34%. Despite pre-existing associations with benthic cover, all fish groups examined (pomacentrids, labrids, chaetodontids, and acanthurids) were resistent to benthic disturbances at the level and scale at which they were applied. Partial Mantel's tests, in combination with partial Canonical Correspondence Analysis enabled spatial and temporal variation to be factored out from experimental effects. Most of the variation in the fish community could be assigned to spatio-temporal variables, indicating that spatial structure over the reef landscape may moderate localised disturbance effects. This study indicates that coral reef fish assemblages may be more resistant to disturbance than many correlative studies would suggest, and highlights a need for further information on levels and scales of natural habitat disturbance in order to apply a structured approach to the experimental investigation of the importance of habitat in structuring coral-reef fish assemblages.