Economies of scaling: More evidence that allometry of metabolism is linked to activity, metabolic rate and habitat

Organismal metabolic rates influence many ecological processes, and the mass-specific metabolic rate of organisms decreases with increasing body mass according to a power law. The exponent in this equation is commonly thought to be the three-quarter-power of body mass, determined by fundamental physical laws that extend across taxa. However, recent work has cast doubt as to the universality of this relationship, the value of 0.75 being an interspecies 'average' of scaling exponents that vary naturally between certain boundaries. There is growing evidence that metabolic scaling varies significantly between even closely related species, and that different values can be associated with lifestyle, activity and metabolic rates. Here we show that the value of the metabolic scaling exponent varies within a group of marine ectotherms, chitons (Mollusca: Polyplacophora: Mopaliidae), and that differences in the scaling relationship may be linked to species-specific adaptations to different but overlapping microhabitats. Oxygen consumption rates of six closely related, co-occurring chiton species from the eastern Pacific (Vancouver Island, British Columbia) were examined under controlled experimental conditions. Results show that the scaling exponent varies between species (between 0.64 and 0.91). Different activity levels, metabolic rates and lifestyle may explain this variation. The interspecific scaling exponent in these data is not significantly different from the archetypal 0.75 value, even though five out of six species-specific values are significantly different from that value. Our data suggest that studies using commonly accepted values such as 0.75 derived from theoretical models to extrapolate metabolic data of species to population or community levels should consider the likely variation in exponents that exists in the real world, or seek to encompass such error in their models. This study, as in numerous previous ones, demonstrates that scaling exponents show large, naturally occurring variation, and provides more evidence against the existence of a universal scaling law. © 2012 Elsevier B.V.

Does wave exposure determine the interactive effects of losing key grazers and ecosystem engineers?

The consequences of biodiversity loss in the face of environmental change remain difficult to predict, given the complexity of interactions among species and the context-dependency of their functional roles within ecosystems. Predictions may be enhanced by studies testing how the interactive effects of species loss from different functional groups vary with important environmental drivers. On rocky shores, limpets and barnacles are recognised as key grazers and ecosystem engineers, respectively. Despite the large body of research examining the combined effects of limpet and barnacle removal, it is unclear how their relative importance varies according to wave exposure, which is a dominant force structuring intertidal communities. We tested the responses of algal communities to the removal of limpets and barnacles on three sheltered and three wave-exposed rocky shores on the north coast of Ireland. Limpet removal resulted in a relative increase in microalgal biomass on a single sheltered shore only, but led to the enhanced accumulation of ephemeral macroalgae on two sheltered shores and one exposed shore. On average, independently of wave exposure or shore, ephemeral macroalgae increased in response to limpet removal, but only when barnacles were removed. On two sheltered shores and one exposed shore, however, barnacles facilitated the establishment of fucoid macroalgae following limpet removal. Therefore, at the scale of this study, variability among individual shores was more important than wave exposure per se in determining the effect of limpet removal and its interaction with that of barnacles. Overall, these findings demonstrate that the interactive effects of losing key species from different functional groups may not vary predictably according to dominant environmental factors.

The influence of water motion on the growth rate of the kelp Laminaria hyperborea

The kelp Laminaria hyperborea is a dominant component of the subtidal nearshore ecosystem and is subjected to a heterogeneous wave and current climate. Water motion is known to influence physiological processes in macroalgae such as photosynthesis and nutrient uptake attributed to mass-transfer limitation. The study attempts to establish the effect of water motion on the growth rates of blades and elongation rates of the stipes of L. hyperborea at adjacent wave-exposed and wave-sheltered locations over a 12month period from field observations. The observations were supported by detailed physical and chemical measurements (light, temperature, seawater nutrient concentrations and hydrodynamics) and of tissue carbon and nitrogen concentrations together with δ¹³carbon. Despite a 30% difference in the root mean square of the velocity (Vel_rms) between the two survey locations, there was no evidence to suggest that water motion had any direct influence on the growth rates of either the blades or elongation of stipes of L. hyperborea. No significant differences were observed between either environmental or plant physiological variables between the sheltered and exposed locations. Using an integral velocity parameter (Vel_rms) the present study also highlighted the importance of the tidally induced current component of water flow in the subtidal zone. 

One size fits all: stability of metabolic scaling under warming and ocean acidification in echinoderms

Responses by marine species to ocean acidification (OA) have recently been shown to be modulated by external factors including temperature, food supply and salinity. However the role of a fundamental biological parameter relevant to all organisms, that of body size, in governing responses to multiple stressors has been almost entirely overlooked. Recent consensus suggests allometric scaling of metabolism with body size differs between species, the commonly cited 'universal' mass scaling exponent (b) of A3/4 representing an average of exponents that naturally vary. One model, the Metabolic-Level Boundaries hypothesis, provides a testable prediction: that b will decrease within species under increasing temperature. However, no previous studies have examined how metabolic scaling may be directly affected by OA. We acclimated a wide body-mass range of three common NE Atlantic echinoderms (the sea star Asterias rubens, the brittlestars Ophiothrix fragilis and Amphiura filiformis) to two levels of pCO(2) and three temperatures, and metabolic rates were determined using closed-chamber respirometry. The results show that contrary to some models these echinoderm species possess a notable degree of stability in metabolic scaling under different abiotic conditions; the mass scaling exponent (b) varied in value between species, but not within species under different conditions. Additionally, we found no effect of OA on metabolic rates in any species. These data suggest responses to abiotic stressors are not modulated by body size in these species, as reflected in the stability of the metabolic scaling relationship. Such equivalence in response across ontogenetic size ranges has important implications for the stability of ecological food webs.

Global mitochondrial DNA phylogeography and population structure of the silky shark, Carcharhinus falciformis

Globally, sharks are under enormous pressure from fishing efforts. One such species is the silky shark, Carcharhinus falciformis, which occurs in all the Earth’s tropical oceans and is captured in large numbers in pelagic fisheries. Regionally, the silky shark is listed as Vulnerable to Near Threatened by the International Union for the Conservation of Nature due to high levels of direct and by catch exploitation. Despite major conservation concerns about this species, little is known about its genetic status and level of demographic or evolutionary connectivity among its regional distributions. We report a genetic assessment of silky sharks sampled across a major portion of the species’ global range. We sequenced the complete mitochondrial DNA control region from 276 individuals taken from the western Atlantic and Indo-Pacific Oceans and the Red Sea. Overall, haplotype and nucleotide diversities were relatively large (0.93 ± 0.01 and 0.61 ± 0.32 %, respectively). Nucleotide diversity in Indo-Pacific sharks, however, was significantly lower and about half that in Atlantic sharks. Strong phylogeographic partitioning occurred between ocean basins. Furthermore, shallow but significant pairwise statistical differentiation occurred among most regional samples within the Indo-Pacific, but not the western Atlantic. Overall, at least five mitochondrial DNA populations of silky sharks were identified globally. Despite historically large population sizes, silky sharks appear to be isolated on relatively small spatial scales, at least in the Indo-Pacific, indicating that conservation and management efforts will need to be exerted at relatively small scales in a pelagic and highly vagile species.

Phylogeographical analyses of shellfish viruses: inferring a geographical origin for ostreid herpesviruses OsHV-1 (Malacoherpesviridae)

Mortality episodes have regularly been affecting the shellfish industry throughout its history. Some of these mortalities, especially in the oyster industry, have been attributed to herpesviruses. Purification of viral particles and molecular characterization have led to the development of routine monitoring, as well as improved taxonomic classification. Ostreid herpesviruses (Malacoherpesviridae), mostly affecting Pacific oysters Crassostrea gigas, have been sporadically recorded in the French oyster industry since the early 1990s (OsHV-1 'reference'). From 2008, a new variant of ostreid herpesvirus (OsHV-1 mu Var) has emerged and seriously impacted oyster production in France and other European countries. Consequently, the presence of ostreid herpesviruses has been monitored in different oyster producing areas around the world. The present study compiles molecular data that are available from survey efforts and takes a biogeographical approach, in order to infer an origin for ostreid herpesviruses. The highest genotype diversity was found in East Asia, despite a lower survey effort in that area than in Europe. Genotype network analyses show that both populations of ostreid herpesviruses present in Europe (OsHV-1 'reference' and OsHV-1 mu Var) are closely related to genotypes recorded in Asia. Moreover, ostreid herpesviruses have been detected in wild and symptom-free populations of various Asian native Crassostrea species. In the rest of the world, ostreid herpesvirus genotypes were recorded from cultivated C. gigas, and mostly associated with mortality episodes. Results of this study are therefore highly suggestive of an Asian origin for these viruses, which can be pathogenic under farming conditions. It also highlights the risks of European stock improvements, by means of overseas shellfish imports.

Forecasting invasions: resource use by mussels informs invasion patterns along the South African coast

Invasive species are often more able to rapidly and efficiently utilise resources than natives, and comparing per capita resource use at different resource densities among invaders and trophically analogous natives could allow for reliable predictions of invasiveness. In South Africa, invasion by the Mediterranean mussel Mytilus galloprovincialis has transformed wave-exposed shores, negatively affecting native mussel species. Currently, South Africa is experiencing a second mussel invasion with the recent detection of the South American Semimytilus algosus. We tested per capita uptake of an algal resource by invading M. galloprovincialis, S. algosus, and the native Aulacomya atra at different algal concentrations and temperatures, representing the west and south coasts of South Africa, to examine whether their per capita resource use could be a predictor of their spread and subsequent invasiveness. Regardless of temperature, M. galloprovincialis was the most efficient consumer, significantly reducing algal cells compared to the other species when the resource was presented in both low and high starting densities. Furthermore, these findings aligned with a greater biomass of M. galloprovincialis on the shore in comparison with the other species. Resource use by the new invader S. algosus was dependent on the density of resource and, although this species was efficient at low algal concentrations at cooler temperatures, this pattern broke down at higher algal densities. This was once more reflected in lower biomass in surveys of this species along the cool west coast. We therefore forecast that S. algosus will be become established along the south coast; however, we also predict that M. galloprovincialis will maintain dominance on these shores.

Footprints in the sand: a persistent spatial impression of fishing in a mobile groundfish assemblage

Fishing is well known to curtail the size distribution of fish populations. This paper reports the discovery of small-scale spatial patterns in length appearing in several exploited species of Celtic Sea demersal 'groundfish'. These patterns match well with spatial distributions of fishing activity, estimated from vessel monitoring records taken over a period of 6 years, suggesting that this 'mobile' fish community retains a persistent impression of local-scale fishing pressure. An individual random-walk model of fish movement best matched these exploitation 'footprints' with individual movement rates set to <35 km per year. We propose that Celtic Sea groundfish may have surprisingly low movement rates for much of the year, such that fishing impact is spatially heterogeneous and related to local fishing intensity.

Patterns of co‑occurrence and interactions between age classes of the common triplefin, Forsterygion lapillum

Many marine organisms have pelagic larval stages that settle into benthic habitats occupied by older individuals; however, a mechanistic understanding of inter cohort interactions remains elusive for most species. Patterns of spatial covariation in the densities of juvenile and adult age classes of a small temperate reef fish, the common triplefin (Forsterygion lapillum), were evaluated during the recruitment season (Feb–Mar, 2011) in Wellington, New Zealand (41°17′S, 174°46′E). The relationship between juvenile and adult density among sites was best approximated by a dome-shaped curve, with a negative correlation between densities of juveniles and adults at higher adult densities. The curve shape was temporally variable, but was unaffected by settlement habitat type (algal species). A laboratory experiment using a “multiple-predator effects”design tested the hypothesis that increased settler mortality in the presence of adults (via enhanced predation risk or cannibalism) contributed to the observed negative relationship between juveniles and adults. Settler mortality did not differ between controls and treatments that contained either one (p = 0.08) or two (p = 0.09) adults. However, post hoca analyses revealed a significant positive correlation between the mean length of juveniles used in experimental trials and survival of juveniles in these treatments, suggesting that smaller juveniles may be vulnerable to cannibalism. There was no evidence for risk enhancement or predator interference when adults were present alongside a hetero specific predator (F. varium). These results highlight the complex nature of intercohort relationships in shaping recruitment patterns and add to the growing body of literature recognizing the importance of age class interactions.

The influence of water motion on the growth rate of the kelp Laminaria digitata

The shallow water kelp Laminaria digitata, abundant in coastal zones of the North Atlantic, is exposed to a range of hydrodynamic environments that makes it ideal for assessing the role of water motion on their growth rate. Here we quantify the growth of L. digitata, as a factor of blade and stipe elongation, at sites adjacent to Strangford Lough, Northern Ireland under different hydrodynamic conditions over a one year period. A modelling approach was used to numerically determine both the temporal and spatial variability of the hydrodynamic environment. Ambient seawater nutrient concentrations, temperature and irradiance were measured as well as the internal nutrient status of the L. digitata populations. Kelp populations growing in the greatest and lowest water motion showed the lowest growth rates. Differences observed in growth rate could not be attributed to seawater nutrient availability, temperature or light. The internal nutrient status also suggested no influence on the observed differences in growth rate. Therefore if there are minimal differences in light, temperature and nutrients between sites, then populations of L. digitata exposed to different water motions are likely to exhibit different growth rates. It is suggested that the growth rate differences observed were a function of water motion with the possibility that, in response to the hydrodynamic forces experienced by the algal cells, L. digitata kelps in the high energy environments were putting more energy into strengthening cell walls rather than blade elongation

Validating the use of intrinsic markers in body feathers to identify inter‑individual differences in non‑breeding areas of northern fulmars

Many wildlife studies use chemical analyses to explore spatio-temporal variation in diet, migratory patterns and contaminant exposure. Intrinsic markers are particularly valuable for studying non-breeding marine predators, when direct methods of investigation are rarely feasible. However, any inferences regarding foraging ecology are dependent upon the time scale over which tissues such as feathers are formed. In this study, we validate the use of body feathers for studying non-breeding foraging patterns in a pelagic seabird, the northern fulmar. Analysis of carcasses of successfully breeding adult fulmars indicated that body feathers moulted between September and March, whereas analyses of carcasses and activity patterns suggested that wing feather and tail feather moult occurred during more restricted periods (September to October and September to January, respectively). By randomly sampling relevant body feathers, average values for individual birds were shown to be consistent. We also integrated chemical analyses of body feather with geolocation tracking data to demonstrate that analyses of δ13C and δ15N values successfully assigned 88 % of birds to one of two broad wintering regions used by breeding adult fulmars from a Scottish study colony. These data provide strong support for the use of body feathers as a tool for exploring non-breeding foraging patterns and diet in wide-ranging, pelagic seabirds.