What is hidden behind the concept of ecosystem efficiency in energy transformation?

The number of energy transformation levels in trophic webs is usually below five, but can be extended up to ten when parasites and hyper-parasites are included. Research on the structure and function of food webs is relevant to the complexity–stability–productivity debate. The aim of this theoretical analysis is to link energetic and connectional aspects of ecosystems with information theory. Updating an energetic model reported by Ricklefs [Ecologia, Zanichelli Editore S.p.A., Bologna, Italy, 1993, p. 896], our approach is integrated with a static analysis of food webs. The length of food webs is theoretically associated with the average ecological efficiency which can be empirically correlated with the effective connectance between species. Furthermore, the advantage of greater complexity when applied to a signalling network is qualitatively addressed.The overall efficiency of energy transformation into biomass throughout a trophic web, in an ecosystem with a given number of species, is the resultant of the various ecological efficiencies, η, at the transitions between the trophic levels. However, we propose that an increment in effective connectance and interspecies connectivity based on a superimposed signalling web may increase the η values, despite the fact that signalling per se has an energetic cost. According to this hypothesis, ecosystem stability would not be necessarily reduced by increasing the number of trophic levels, N, whenever stability in terms of persistence is improved by a cost-efficient regulatory network.     

The top-down mechanism for body-mass-abundance scaling

Scaling relationships between mean body masses and abundances of species in multitrophic communities continue to be a subject of intense research and debate. The top-down mechanism explored in this paper explains the frequently observed inverse linear relationship between body mass and abundance (i.e., constant biomass) in terms of a balancing of resource biomasses by behaviorally and evolutionarily adapting foragers, and the evolutionary response of resources to this foraging pressure. The mechanism is tested using an allometric, multitrophic community model with a complex food web structure. It is a statistical model describing the evolutionary and population dynamics of tens to hundreds of species in a uniform way. Particularities of the model are the detailed representation of the evolution and interaction of trophic traits to reproduce topological food web patterns, prey switching behavior modeled after experimental observations, and the evolutionary adaptation of attack rates. Model structure and design are discussed. For model states comparable to natural communities, we find that (1) the body-mass abundance scaling does not depend on the allometric scaling exponent of physiological rates in the form expected from the energetic equivalence rule or other bottom-up theories; (2) the scaling exponent of abundance as a function of body mass is approximately -1, independent of the allometric exponent for physiological rates assumed; (3) removal of top-down control destroys this pattern, and energetic equivalence is recovered. We conclude that the top-down mechanism is active in the model, and that it is a viable alternative to bottom-up mechanisms for controlling body-mass-abundance relations in natural communities.     

Biochemical and energetic composition of bathyal echinoids and an asteroid,holothuroid and crinoid from the Bahamas

The biochemical and energetic composition of body components of ten species of bathyal echinoids, and an asteroid, a holothuroid and a stalked crinoid were determined from individuals sampled from a variety of deep-water sites near the Bahamas (north Caribbean Sea) in October 1988. When compared with other studies of echinoderms, no geographic- or depth-related differences in biochemical or energetic composition were found. Body-wall tissues were composed primarily of skeletal material (mineral ash), but were comparatively high in organic material in the echinothuriid echinoids, and the asteroid and holothuroid. Gut tissues and pyloric cecae had high levels of lipid and protein, indicating their potential role in nutrient storage. Body-wall tissues were generally low in energy, but were highest in the echinoidsAraeosoma belli (7.7 kJ g^–1 dry wt) andSperosoma antillense (8.0 kJ g^–1 dry wt), the asteroidOphidiaster alexandri (8.9 kJ g^–1 dry wt), and the holothuroidEostichopus regalis (13.1 kJ g^–1 dry wt). Energy levels of gut and pyloric cecal tissues were two to three times higher than those of body-wall tissues. Total somatic tissue energy values varied greatly among species, ranging from 1.5 kJ in the echinoidAspidodiadema jacobyi to 142.1 kJ inE. regalis. As the bathyal echinoderms examined in this study occur in great abundance, they represent a significant reservoir of organic and inorganic materials and energy in deep-water benthic systems.     

Free energy flow in the process of collecting material for body build up of aquatic organisms

A general formula defining the free energy expenditures in the processes of algal nutrition and zooplankton feeding has been derived. It appears to be symmetrical in respect of space and time scales. In conditions of optimal control the theoretical relationship between effective free energy expenditures and losses appears to be the ‘golden proportion’. The general trends of the system due to the first and second laws of thermodynamics and Prigogine's theorem form a theoretical basis for phenomenological rules of ecosystem's development.     

Feasibility and stability in randomly assembled Lotka-Volterra models

For a Lotka-Volterra model to represent a viable ecosystem it's nontrivial equilibrium must be feasible. If m is the number of species, it is shown that in a set of randomly assembled Lotka-Volterra models, the fraction of models with a feasible equilibrium is some function of m which behaves like 2^?m. Moreover a subset of Lotka-Volterra models, each of which has a feasible equilibrium, has the same stability property as a set of linear models which is assembled randomly in the same manner. This contradicts a recent claim that a Lotka-Volterra model with a feasible equilibrium tends to be stable. Thus for two reasons the probability that a Lotka-Volterra model represents a viable and stable ecosystem decreases rapidly with the number of species. This supports the theme developed by May that stability in model ecosystems decreases with diversity.     

Variability of epipelagic macrozooplankton/micronekton community structure in the NW Weddell Sea,Antarctica (1995–1996)

Macrozooplankton/micronekton in the epipelagic zone (0–100 m) of the NW Weddell Sea were sampled with a 10 m² multiple opening-closing net environmental sampling system (MOCNESS) trawl during three cruises in 1995 and 1996. A total of 40 species were collected during all cruises. Community composition, as measured using Bray-Curtis analysis and multi-dimensional scaling, permitted separation into two groups: group 1 included samples collected during September–October 1995 and group 2 contained samples collected during April-May 1996 and November–December 1996. Zooplankton collections were dominated by Thysanoessa macrura in September–October 1995 (high pack-ice cover) and by both Euphausia superba and Salpa thompsoni in April–May 1996 (intermediate pack-ice cover) and November–December 1996 (open water). Copepods were not sampled quantitatively by the MOCNESS trawl and were not included in this analysis. Trends in measured parameters of community structure are discussed with respect to environmental characteristics during each sampling period.Communicated by P.W. Sammarco, Chauvin     

Phylogeny and metabolic scaling in mammals

The scaling of metabolic rates to body size is widely considered to be of great biological and ecological importance, and much attention has been devoted to determining its theoretical and empirical value. Most debate centers on whether the underlying power law describing metabolic rates is 2/3 (as predicted by scaling of surface area/volume relationships) or 3/4 ("Kleiber's law"). Although recent evidence suggests that empirically derived exponents vary among clades with radically different metabolic strategies, such as ectotherms and endotherms, models, such as the metabolic theory of ecology, depend on the assumption that there is at least a predominant, if not universal, metabolic scaling exponent. Most analyses claimed to support the predictions of general models, however, failed to control for phylogeny. We used phylogenetic generalized least-squares models to estimate allometric slopes for both basal metabolic rate (BMR) and field metabolic rate (FMR) in mammals. Metabolic rate scaling conformed to no single theoretical prediction, but varied significantly among phylogenetic lineages. In some lineages we found a 3/4 exponent, in others a 2/3 exponent, and in yet others exponents differed significantly from both theoretical values. Analysis of the phylogenetic signal in the data indicated that the assumptions of neither species-level analysis nor independent contrasts were met. Analyses that assumed no phylogenetic signal in the data (species-level analysis) or a strong phylogenetic signal (independent contrasts), therefore, returned estimates of allometric slopes that were erroneous in 30% and 50% of cases, respectively. Hence, quantitative estimation of the phylogenetic signal is essential for determining scaling exponents. The lack of evidence for a predominant scaling exponent in these analyses suggests that general models of metabolic scaling, and macro-ecological theories that depend on them, have little explanatory power.     

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 ¾ 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 pCO₂ 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.     

Scaling toxicity data across species

The response of various species to doses of chemicals can often give the impression that some (such as cattle in the case of molybdenum) are much more susceptible than others to these chemicals. These impressions usually rely on an underlying assumption that equivalent doses are based on mg of the chemical per kg body weight of the animal. That is, that doses scale as the first power of body weight. This assumption is more often wrong than right. When viewed in a more general way, where the scaling is proportional to a power of the body weight and the exponent determined empirically, it is often found that equivalent doses scale with an exponent in the range of 0.6 to 0.8. As a result, larger animals are indeed more susceptible to toxicity on a mg kg^–1 body weight basis, but this is not because of unique differences in the species, but only because of different body sizes. This method of scaling is called allometry or allometric scaling. An early version of this approach was based on body surface area where the exponent is 2/3. More recently, pharmacokinetics has revealed that the reason for the different response of larger animals is related to the slower metabolic and clearance rates for larger animals which give rise to larger biological half-lives for chemicals in the body and to higher tissue concentrations per given dose.     

Exploring community assembly through an individual-based model for trophic interactions

Traditionally, the dynamics of community assembly has been analyzed by means of deterministic models of differential equations. Despite the theoretical advances provided by such models, they are restricted to questions about community-wide features. The individual-based modeling offers an opportunity to link bionomic features to patterns at the community scale, allowing us to understand how trait-based assembly rules can arise by dynamical processes. The present paper introduces an individual-based model of community assembly, and discusses some of the major advantages and drawbacks of this approach. The model was framed to deal with predation among size-structured populations, incorporating allometric constraints to energetic requirements, movement, life-history features and interaction relationships among individuals. A protocol of assembly procedure is proposed, in which a period of intense species introductions is followed by a period without introductions. The resultant communities did not present any pattern of trait over-dispersion, meaning that the multivariate distances of bionomic features among co-occurring species were neither larger nor more regular than expected in a random collection of species. It suggests a weak influence of interspecific interactions in the model environment and individualistic rules of coexistence, driven mainly by the spatial structure. This highlights that trait over-dispersion and resource partitioning should not be considered a necessary condition for coexistence, even in communities entirely structured by internal processes like predation and competition.     

Labor market estimates of the senior discount for the value of statistical life

This article develops the first measures of age–industry job risks to examine the age variations in the value of statistical life. Because of the greater risk vulnerability of older workers, they face flatter wage-risk gradients than younger workers, which we show to be the case empirically. Accounting for this heterogeneity in hedonic market equilibria leads to estimates of the value of statistical life–age relationship that follows an inverted U shape. The estimates of the value of statistical life range from $6.4 million for younger workers to a peak of $9.0 million for those aged 35–44, and then a decline to $3.8 million for those aged 55–62. The decline of the estimated value of statistical life with age is consistent with there being some senior discount in the Clear Skies Initiative analysis.     

Statistical selection of perimeter-area models for patch mosaics in multiscale landscape analysis

This paper presents a statistical method for detecting distinct scales of pattern for mosaics of irregular patches, by means of perimeter–area relationships. Krummel et al. (1987) were the first to develop a method for detecting different scaling domains in a landscape of irregular patches, but this method requires investigator judgment and is not completely satisfying. Grossi et al. (2001) suggested a modification of Krummel's method in order to detect objectively the change points between different scaling domains. Their procedure is based on the selection of the best piecewise linear regression model using a set of statistical tests. Even though the change points were estimated, the null distributions used for testing purposes were those appropriate for known change points. The present paper investigates the effect that estimating the change points has on the underlying distribution theory. The procedure we suggest is based on the selection of the best piecewise linear regression model using a likelihood ratio (LR) test. Each segment of the piecewise linear model corresponds to a fractal domain. Breakpoints between different segments are unknown, so the piecewise linear models are non-linear. In this case, the frequency distribution of the LR statistic cannot be approximated by a chi-squared distribution. Instead, Monte Carlo simulation is used to obtain an empirical null distribution of the LR statistic. The suggested method is applied to three patch types (CORINE biotopes) located in the Val Baganza watershed of Italy.     

Metabolism and swimming efficiency of the bonnethead sharkSphyrna tiburo

Routine metabolic rates of bonnethead sharks,Sphyrna tiburo, of 95 to 4 650 g, ranged from 70.4 to 15.0 kcal kg^–1 d^–1. Over the size range 34 to 95 cm total length, shark swimming-velocities varied from about 29 to 67 cm s^–1. Swimming velocities predicted using Weih's cost-optimization model were similar to observed velocities. The total cost of transport (the energetic cost of transporting 1 unit of body mass 1 km distance) for 1 to 8 kg sharks varied from 0.67 to 0.40 cal g^–1 km^–1. The energetic range (an estimation of the distance traveled after a 25% reduction in body weight) indicates that a 1 kg bonnethead shark would travel 500 km distance in 17 d before displaying a 25% reduction in weight. An 8 kg individual would travel 830 km in 23 d. Although the bonnethead shark is a continuously active species, its routine metabolic rate and the efficiency of its locomotory system may be similar to that of typical bony fishes.     

Assessment of the contribution of land use pattern and intensity to landscape quality: use of a landscape indicator

The objective of this research work is the evaluation of the impact of landuse pattern and intensity on landscape by means of an indicator. The method used to calculate a ‘landscape indicator’ (I_land) allows to take into account the objective as well as the subjective approach of landscape. I_land corresponds to the degree of agreement between landscape supply by farmers and landscape demand by the social groups. The supply and the demand are evaluated through four criteria: ‘diversity’, ‘upkeep’, ‘openness’ and ‘heritage’. The landscape supply is calculated from data of landscape objects (punctual, linear and spatial) for each criterion recorded at the field level. The values of the four criteria for the landscape demand are allocated by the user(s) of the indicator (decision makers, regional council, social groups…) into five classes (0–4). The value of the landscape indicator is the least favourable difference between supply and demand for the four criteria. An example of calculation of the ‘landscape indicator’ for an arable farm is given. The collection of data needs 2 h with the farmer and 2 h for a survey of the farm land.     

The evolution of eusociality in molerats (Bathyergidae): a question of risks,numbers, and costs

Summary This paper addresses risk and energetic considerations fundamentaly causative in the evolution of eusociality in the bathyergid molerats. Three simple mathematical models are presented which predict the probability of successful outbreeding in terms of dispersal risks and the energetic costs of foraging. The predictions of the models are compared with data from the literature, which mostly provide excellent empirical and theoretical support.Inter-habitat dispersal risks are influenced most importantly by group size and resource characteristics, but also by body size, metabolic rate, thermoregulation, soil conditions, and caste structure. Intea-habitat foraging risks are temporaly dependent on rainfall — a factor critical for appropriate dispersal timing. High dispersal and foraging risks necessitate large group sizes and preclude a solitary existence. Outbreeding or inbreeding options are strongly influenced by dispersal risks, with high genetic relatedness in high risk habitats the likely consequence. Offspring should attain inclusive fitness values equal, if not more, than those possible by outbreeding by staying and helping with the colony reproductive effort.Offprint requests to: B.G. Love grove at the German address     

Does begging affect growth in nestling tree swallows,<Emphasis Type="Italic"> Tachycineta bicolor</Emphasis>?

Much of the theoretical work on the evolution of begging assumes this elaborate display is costly. The evidence for an energetic cost to begging has, however, been equivocal. Metabolic studies on nestling birds suggest that begging requires minimal energy, but some growth studies have shown that excess begging reduces growth rates. One difficulty in interpreting these results is that metabolic and growth studies have each been performed on different species. Here, we test whether high begging frequencies depress growth in nestling tree swallows, Tachycineta bicolor, a species in which the metabolic cost of begging has been measured. When we compared the growth of nestlings stimulated to beg at either high or low frequencies, we found no significant differences in their mass gained, wing growth or portion of ingested energy devoted to begging either during the experimental period or in the 24 h following the end of the experiment. We also found no significant relationship between begging intensity and growth measurements. The results of our study are consistent with previous metabolic studies on this species suggesting that the energetic cost of begging is relatively low. More generally, evidence for a fitness cost of begging via decreased growth is equivocal.Communicated by J. Dickinson     

Transient nestmate recognition cues contribute to a multicolonial population structure in the ant,Leptothorax curvispinosus

Summary Individual nests of the facultatively polygynous and polydomous ant, Leptothorax curvispinosus, were mapped at two sites, collected, and maintained under uniform laboratory conditions. Tests of worker acceptability between nests were conducted 2–4 weeks and 13–17 weeks after collection. Nests collected near to one another (0.09–1.87 m) were sometimes nonaggressive, and were significantly less aggressive than those from different sites (7 km apart); and there was no significant difference in aggressiveness between tests for these distance categories. However, aggression between nests collected farther away from one another at the same site (1.52–4.65 m) decreased significantly between tests: initially, the level of aggressiveness was equivalent to that between nests from different sites but later it was reduced to that between near nests. These results indicate that polydomous colonies of this species occur within multicolonial populations; and that colony segregation within local populations is largely maintained by transient environmentally-based nestmate recognition cues. More stable cues of genetic or environmental origin (or both) are also present and contribute to discrimination even after extended periods of culture under uniform conditions. These results suggest that the maintenance of colony autonomy within genetically highly interrelated populations may be the prime function of environmentally-based nestmate recognition cues. Colony autonomy under such circumstances may be important to maintain a relatively small but optimal colony size, or because the mechanisms which regulate colony growth, development, etc., require a limited colony size.     

Dynamic environ analysis of compartmental systems: A computational approach

Ecosystems are often modeled as stocks of matter or energy connected by flows. Network environ analysis (NEA) is a set of mathematical methods for using powers of matrices to trace energy and material flows through such models. NEA has revealed several interesting properties of flow–storage networks, including dominance of indirect effects and the tendency for networks to create mutually positive interactions between species. However, the applicability of NEA is greatly limited by the fact that it can only be applied to models at constant steady states. In this paper, we present a new, computationally oriented approach to environ analysis called dynamic environ approximation (DEA). As a test of DEA, we use it to compute compartment throughflow in two implementations of a model of energy flow through an oyster reef ecosystem. We use a newly derived equation to compute model throughflow and compare its output to that of DEA. We find that DEA approximates the exact results given by this equation quite closely – in this particular case, with a mean Euclidean error ranging between 0.0008 and 0.21 – which gives a sense of how closely it reproduces other NEA-related quantities that cannot be exactly computed and discuss how to reduce this error. An application to calculating indirect flows in ecosystems is also discussed and dominance of indirect effects in a nonlinear model is demonstrated.     

Polychaetes associated to a<Emphasis Type="Italic"> Cymodocea nodosa</Emphasis> meadow in the Canary Islands: assemblage structure,temporal variability and vertical distribution compared to other Mediterranean seagrass meadows

The structure, diversity and temporal distribution of the infaunal polychaetes associated with Cymodocea nodosa meadows were studied in Tenerife (Canary Islands). The samples were collected monthly throughout a year, to depths of 13–16 m. The sediment was extracted by means of PVC cores, in which four layers were separated (i.e. 0–5 cm, 5–10 cm, 10–20 cm and 20–30 cm). A total of 1,167 polychaete specimens, belonging to 69 taxa were collected, representing one of the most dominant groups in the benthic assemblage throughout the entire year. The most common families were Syllidae, Paraonidae and Spionidae, both in terms of abundance and species richness. The dominant species were Streptosyllis bidentata, Aricidea assimilis and Exogone parahomoseta mediterranea, representing also the only constant species throughout the year. The highest values of species richness, diversity, equitability and abundance of polychaetes occurred in September. The multifactorial analysis of abundances (i.e. cluster analysis and non-metric, multi-dimensional scaling) indicated temporal segregation of the samples from July, August and September (i.e. the warmest months) with respect to those from the rest of the year, due to structural differences in the assemblage. Polychaete species have been found to a depth of up to 30 cm in the sediment. Nevertheless, most of them (89%) occurred in the upper 5 cm of the sediment, with an increase of specimens in deeper layers in February (i.e. due to sporadic episodes of higher hydrodynamics). To compare the vertical distribution of polychaetes, additional core samples were collected in two seagrass meadows (i.e. C. nodosa and Ruppia cirrhosa) at Ebros Delta (NW Mediterranean); these were separated into five layers (i.e. 0–5 cm, 5–10 cm, 10–15 cm, 15–20 cm, 20–25 cm). The results obtained for the R. cirrhosa meadow (98% of the polychaetes within the upper 5 cm) agree with those for the Canarian C. nodosa meadow, while the polychaetes reached up to 15 cm depth in the Mediterranean C. nodosa meadow (i.e. ~39% between 0 and 5 cm, ~41% between 5 and 10 cm, ~20% between 10 and 15 cm). Our results indicated that the structural characteristics of the assemblages appeared to be more strongly controlled by the combined characteristics of the sediment (i.e. lack of oxygen, granulometry and degree of compaction) than by the seagrass species building the meadow.Communicated by O. Kinne, Oldendorf/Luhe