Spatial variance in abundance and occupancy of corals across broad geographic scales

Species assemblages vary in structure due to a wide variety of processes operating at ecological and much broader biogeographical scales. Cross-scale studies of assemblage structure are necessary to fully understand this variability. Here, we evaluate the abundance and occupancy patterns of hierarchically sampled coral assemblages in three habitats (reef flat, crest, and slope) and five regions (Indonesia, Papua New Guinea, the Solomon Islands, American Samoa, and the Society Islands) across the west-central Pacific Ocean. Specifically, we compare two alternative models that unify spatial variance and occupancy via the negative binomial distribution. The first assumes a power-law scaling between the mean and variance of abundance; the second assumes a quadratic variance-mean relationship and a constant abundance-invariant aggregation parameter. Surprisingly, the well-established power-law model performs worse than the model assuming abundance-invariant aggregation, for both variance-mean and occupancy-abundance relationships. We also find strong evidence for regional and habitat variation in these relationships and in the levels of aggregation estimated by the abundance-invariant aggregation model. Among habitats, corals on reef flats exhibited lower occupancy and higher levels of aggregation compared to reef crests and slopes. Among regions, low occupancy and high aggregation were most pronounced across all habitats in American Samoa. These patterns may be related to habitat and regional differences in disturbance and recovery processes. Our results suggest that the spatial scaling of abundance and occupancy is sensitive to processes operating among these habitats and at regional scales. However, the consistency of these relationships across species within assemblages suggests that a theoretical unification of spatial variance and occupancy patterns is indeed possible.     

Regional variation in the hierarchical partitioning of diversity in coral-dwelling fishes

The size of the regional species pool may influence local patterns of diversity. However, it is unclear whether certain spatial scales are less sensitive to regional influences than others. Additive partitioning was used to separate coral-dwelling fish diversity to its alpha and beta components, at multiple scales, in several regions across the Indo-Pacific. We then examined how the relative contribution of these components changes with increased regional diversity. By employing specific random-placement null models, we overcome methodological problems with local-regional regressions. We show that, although alpha and beta diversities within each region are consistently different from random-placement null models, the increase in beta diversities among regions was similar to that predicted once heterogeneity in coral habitat was accounted for. In contrast, alpha diversity within single coral heads was limited and increased less than predicted by the null models. This was correlated with increased intraspecific aggregation in more diverse regions and is consistent with ecological limitations on the number of coexisting species at the local scale. These results suggest that, apart from very small spatial scales, variation in the partitioning of fish diversity along regional species richness gradients is driven overwhelmingly by the corresponding gradients in coral assemblage structure.     

Risk spreading, connectivity, and optimal reserve spacing

Two important processes determining the dynamics of spatially structured populations are dispersal and the spatial covariance of demographic fluctuations. Spatially explicit approaches to conservation, such as reserve networks, must consider the tension between these two processes and reach a balance between distances near enough to maintain connectivity, but far enough to benefit from risk spreading. Here, we model this trade-off. We show how two measures of metapopulation persistence depend on the shape of the dispersal kernel and the shape of the distance decay in demographic covariance, and we consider the implications of this trade-off for reserve spacing. The relative rates of distance decay in dispersal and demographic covariance determine whether the long-run metapopulation growth rate, and quasi-extinction risk, peak for adjacent patches or intermediately spaced patches; two local maxima in metapopulation persistence are also possible. When dispersal itself fluctuates over time, the trade-off changes. Temporal variation in mean distance that propagules are dispersed (i.e., propagule advection) decreases metapopulation persistence and decreases the likelihood that persistence will peak for adjacent patches. Conversely, variation in diffusion (the extent of random spread around mean dispersal) increases metapopulation persistence overall and causes it to peak at shorter inter-patch distances. Thus, failure to consider temporal variation in dispersal processes increases the risk that reserve spacings will fail to meet the objective of ensuring metapopulation persistence. This study identifies two phenomena that receive relatively little attention in empirical work on reserve spacing, but that can qualitatively change the effectiveness of reserve spacing strategies: (1) the functional form of the distance decay in covariance among patch-specific demographic rates and (2) temporal variation in the shape of the dispersal kernel. The sensitivity of metapopulation recovery and persistence to how covariance of vital rates decreases with distance suggests that estimating the shape of this function is likely to be as important for effective reserve design as estimating connectivity. Similarly, because temporal variation in dispersal dynamics influences the effect of reserve spacing, approaches to reserve design that ignore such variation, and rely instead on long-term average dispersal patterns, are likely to lead to lower metapopulation viability than is actually achievable.     

Setbacks to shoot growth are common in woody plants, so how are shoots of some species safer than others?

Tissue turnover is a critical facet of plant life history variation. This study quantifies losses from setbacks to growth of terminal woody shoots 1.2m long, across 83 species and seven sites in eastern Australia. Setbacks, where the leading meristem had been removed or died and a new leader had emerged, were common (median three per shoot). Shoots had lost an average of 0.25 m of lead-stem length for 1.2 m net shoot-length gain. Insects like girdlers and borers were prominent causes of large setbacks. The sites spanned tropical to temperate and humid to semiarid climates, but variation in stem loss was much greater across species than across sites. We measured 17 plant functional traits related to growth form, mechanics, hydraulics, and economics. Only four traits were correlated with variation across species in stem losses: stem diameter, stem nitrogen content, bark thickness, and maximum photosynthetic rate. The correlations were weak. Stem specific gravity (wood density) showed no correlation with risk. Our results suggest a pattern similar to the growth risk trade-off known for herbaceous plants, where traits associated with fast growth increase tissue turnover and herbivory, but the weak correlations leave ample scope for other influences that remain to be identified.     

Reducing per capita food supply alters urchin condition and habitat

When food supply declines or population density increases, the per capita food availability is reduced causing a decline in condition of consumers. Many consumers alter their feeding behaviour and ultimately the surrounding community (e.g. overgrazing and formation of urchin barrens). This study tested the hypothesis that sea urchin populations are of greater density and poorer condition in barrens (little food) than forest habitat (lots of food). We then tested the hypothesis that a decrease in per capita food supply to urchins has a negative effect not only on their condition but also on their surrounding habitat. We experimentally assessed the effect of limited food supply and increased density of a subtidal Australian sea urchin (Heliocidaris erythrogramma) on their condition (i.e. gonad index) and surrounding benthic habitat (i.e. turf-forming algae). Our results show that a reduction in food supply led to poorer consumer condition and greater herbivory on surrounding local habitat. We provide evidence that per capita food reduction is one of the necessary conditions for the over-consumption by urchins (i.e. urchin barrens), a proposed but previously untested mechanism.     

Effects of four decades of fire manipulation on woody vegetation structure in Savanna

The amount of carbon stored in savannas represents a significant uncertainty in global carbon budgets, primarily because fire causes actual biomass to differ from potential biomass. We analyzed the structural response of woody plants to long-term experimental burning in savannas. The experiment uses a randomized block design to examine fire exclusion and the season and frequency of burn in 192 7-ha experimental plots located in four different savanna ecosystems. Although previous studies would lead us to expect tree density to respond to the fire regime, our results, obtained from four different savanna ecosystems, suggest that the density of woody individuals was unresponsive to fire. The relative dominance of small trees was, however, highly responsive to fire regime. The observed shift in the structure of tree populations has potentially large impacts on the carbon balance. However, the response of tree biomass to fire of the different savannas studied were different, making it difficult to generalize about the extent to which fire can be used to manipulate carbon sequestration in savannas. This study provides evidence that savannas are demographically resilient to fire, but structurally responsive.     

Effects of photoacclimation on the light niche of corals: a process-based approach

The ecology of photosynthetic organisms is influenced by the need to adjust the photosynthetic apparatus to variable light environments (photoacclimation). In this study, we quantified different components of the photoacclimation process for a reef-building coral (Turbinaria mesenterina, Lamarck, 1816): including, variation in absorption cross-section, size of photosynthetic units, turnover time, chlorophyll content, and colony respiration. We used these calibrations to characterize this species’ light niche, and to determine the sensitivity of the niche boundaries to different processes of photoacclimation. Results showed that the breadth of the light niche was most sensitive to the size of the photosynthetic unit, absorption cross-section, and rates of respiration. Habitats with the highest light availability did not lead to maximal energy acquisition. This was because, although corals acclimated to high light have high rates of photosynthesis per unit chlorophyll, their chlorophyll content was strongly reduced. This suggests that potential energetic benefits that could be achieved through increased light harvesting (i.e., increased chlorophyll content) in high-light habitats are outweighed by costs associated with photoprotection. Such costs appear to place an upper bound on the habitat distributions of coral species. Our approach reveals how the photophysiological processes involved in photoacclimation interact to determine the light niche.     

Three-dimensional barricading of a predatory trap reduces predation and enhances prey capture

Animal structures come at material, energetic, time, and expression costs. Some orb-web spiders add three-dimensional barrier structures to their webs, but many do not. Predator protection is considered to be the principal benefit of adding these structures. Accordingly, it remains paradoxical why some orb-web spiders might construct the barriers while others do not. Here, we experimentally determined whether the barrier structure added to the horizontal orb web of the spider Cyrtophora moluccensis deters predators at the cost of reducing the amount of prey captured in the field. We conducted experiments by day and night to assess whether the effects vary with the time of day. We found that the three-dimensional barriers not only offered protection from predatory wasps by day but also enhanced the amount of prey captured by day and night. Moreover, the barrier structure appears particularly useful at catching moths, the largest and most energetically profitable prey that it encounters. We, therefore, concluded that reducing the energetic and time costs associated with producing and depositing extra silk threads is the principal reason why barrier structures are used intermittently among orb-web spiders.     

Matching and mismatching stable isotope (δ13C and δ15N) ratios in fin and muscle tissue among fish species: a critical review

Using non-lethal tissue sampling for stable isotope analysis has become standard in many fields, but not for fishes, despite being desirable when species are rare or protected, when repeated sampling of individuals is required or where removal may bias other analyses. Here, we examine the utility of fish dorsal fin membrane as an alternative to muscle for analyzing δ¹³C and δ¹⁵N ratios in two reef fish species (blue cod Parapercis colias and spotty Notolabrus celidotus) that have differing feeding modes. Both species exhibited evidence of size-based feeding from fin δ¹⁵N values, but not from muscle. Blue cod fin δ¹⁵N increased steadily throughout the sampled size range (213–412 mm fork length), whereas spotty exhibited a distinct ontogenetic diet shift at approximately 120–140 mm fork length after which size-based feeding did not occur. Fin membrane was higher than muscle in δ¹³C in both species and in δ¹⁵N for blue cod, but fin δ¹⁵N was lower than muscle in spotty. The δ¹³C and δ¹⁵N fin–muscle offsets were constant in spotty regardless of size, while in blue cod, δ¹³C was constant with fish size, but δ¹⁵N offsets increased with increasing fish size. Non-lethal sampling utilizing fin tissue can be employed to estimate stable isotope values of muscle in fishes, but it is necessary to assess relationships among tissues and the effects of fish size on isotope values a priori for each species studied. Our data indicated that fin membrane may be a more sensitive tissue than muscle for detecting size-based feeding in some fish species using stable isotopes. A critical literature review revealed inconsistencies in tissue types tested, little understanding of tissue-specific trophic shift or turnover rates, and pseudo-replicated analyses leading to erroneous postulating of 1:1 relationships between tissues.