Population energetics of the intertidal isopod Cirolana harfordi

Laboratory experiments and field measurements generated a population energy-budget estimate for the isopod Cirolana harfordi (Lockington). Assimilation of food energy averaged 88% on a diet of fish. About 35% of assimilated energy is allocated for growth and reproduction, while the bulk of the remainder is used for maintenance and activity (respiration). The high growth efficiency of C. harfordi is discussed with respect to ecological efficiency and energy transfer in marine systems.     

Mitochondrial energetics of benthic and pelagic Antarctic teleosts

Antarctic fauna are highly adapted to the frigid waters of the Southern Ocean. This study describes the in vitro temperature sensitivity of oxygen consumption rates measured in liver mitochondria from the pelagic notothenioid Pleuragramma antarcticum between 5 and 35 °C. Oxygen fluxes were measured after the addition of millimolar levels of pyruvate, malate, succinate and glutamate (state II, LEAK) and saturating levels of ADP [state III, oxidative phosphorylation (OXPHOS)]. State III respiration significantly decreased above 18.7 °C. A comparison of the oxidative capacities among P. antarcticum and other notothenioids showed significant differences in state III respiration, where benthic species exhibited about 50 % lower rates than P. antarcticum. In addition, state III respiration rates normalized per milligram of mitochondrial protein of P. antarcticum were up to eight times higher than state III rates reported in the literature for other notothenioids. The comparatively high respiration rates measured in this study may be explained by our approach, which engaged both complexes I and II under conditions of oxidative phosphorylation. State III rates of independently activated complexes I and II were found to range from 42 to 100 % of rates obtained when both complexes were activated simultaneously in the same species. The remarkable tolerance of P. antarcticum OXPHOS toward warmer temperatures was unexpected for an Antarctic stenotherm and may indicate that thermal sensitivity of their mitochondria is not the driving force behind their stenothermy.     

Community assembly in the presence of disturbance: a microcosm experiment

Ecologists know relatively little about the manner in which disturbance affects the likelihood of alternative community stable states and how the history of community assembly affects the relationship between disturbance and species diversity. Using microbial communities comprising bacterivorous ciliated protists assembled in laboratory microcosms, we experimentally investigated these questions by independently manipulating the intensity of disturbance (in the form of density-independent mortality) and community assembly history (including a control treatment with simultaneous species introduction and five sequential assembly treatments). Species diversity patterns consistent with the intermediate disturbance hypothesis emerged in the controls, as several species showed responses indicative of a tradeoff between competitive ability and ability to recover from disturbance. Species diversity in communities with sequential assembly, however, generally declined with disturbance, owing to the increased extinction risk of later colonizers at the intermediate level of disturbance. Similarities among communities subjected to different assembly histories increased with disturbance, a result due possibly to increasing disturbance reducing the importance of competition and hence priority effects. This finding is most consistent with the idea that increasing disturbance tends to reduce the likelihood of alternative stable states. Collectively, these results indicate the strong interactive effects of disturbance and assembly history on the structure of ecological communities.     

Calling energetics of a neotropical treefrog,Hyla microcephala

Summary We investigated the calling energetics of Hyla microcephala, a neotropical treefrog with an unusually complex vocal repertoire. Males respond to the calls of other individuals by adding secondary click notes to their calls, thereby increasing the total number of notes given per minute. Rates of oxygen consumption of males calling in metabolic chambers were 0.41–2.80 ml O₂/(g·h), corresponding to calling rates of 205–6330 notes/h. Note rate explained 95% of the variance in meta-bolic rate; the effect of variation in body size and temperature was small. Data from playback experiments with males in the field showed that note rate increased as as the number of notes in a stimulus call increased, and this resulted in substantial increases in the cost of calling. Average metabolic rates for males in the field were about 1.7 ml O₂/(g·h), for a net cost of calling of about 20 J/h for an average-size male. However, estimated metabolic rates varied by more than 300% and were strongly influenced by the proximity and calling activity of other males in the chorus. Male H. microcephala appear to conserve energy by reducing calling rates when only a few males are active and increasing calling efforts only when vocal competition among males is intense.     

Influence of food concentration on the physiological energetics and growth ofOstrea edulis larvae

Feeding, respiration and growth rates of oyster (Ostrea edulis L.) larvae reared at five food levels were measured throughout the entire larval period. Energy budgets were derived as a function of alga (Isochrysis galbana Parke) food concentration. Ingestion rate (IR, cells h^-1) and oxygen consumption rate ( , nl h^-1) were almost isometric functions of larval size [ash-free dry weight, (AFDW, g)], characterized by the equations: IR=803.9 AFDW^1.13 and =4.85 AFDW^1.09. Ingested ration was directly correlated to cell concentration up to a maximum at 200 cells l^-1, with further increases failing to support higher ingestion rates. Likewise, growth rate linearly increased with food ration up to 100 cells l^-1 (max. growth efficiency,K ₁=25%) and reached a maximum at 200 cells l^-1 (growth rate=5.6 m d^-1), with further increases in food not supporting significantly faster growth. Maintenance ration was 2 to 3% daily dry weight (DW); optimum ration increased during larval development from 5 to 20% DW; maximum ration was 20% DW. During larval rearing, an increasing feeding schedule of 50, 100 and 200 cells l^-1 from Days 0, 5 and 10, respectively, is recommended.     

Mechanics and energetics of swimming in the small copepodAcanthocyclops robustus (Cyclopoida)

High-speed films of swimmingAcanthocyclops robustus were used to test a crustacean swimming-model based on numerical analysis of thrust production. Predicted body velocities and jump distances were usually within 75% of those observed. Most of the thrust which propels.A. robustus is produced by movements of the 2nd, 3rd, and 4th thoracic swimming legs, with only small contributions from the first thoracic swimming legs. A model analyzed without the first antennae suggested that the antennae do not produce significant thrust. The leg and antennal movements could be described with trigonometric equations (cosine curves), but were best described by polynomial fits of position vs time data from the films. Patterns of swimming velocity varied among four episodes that were modeled, and followed differences in swimming-leg motions. Model results for the small (cephalothorax length = 0.6 mm) cyclopoidA. robustus and those which have been reported for the large calanoid copepodPleuromamma xiphias and other swimmers indicate that mechanical efficiency (30%) does not scale with body size, whereas jump distance (one body length), proportion of thrust generated by hydrodynamic added mass (70%), and net cost of transport,C _p (40 to 109 cal g^–1 km^–1) do.Please address all correspondence and requests for reprints to G. Gust at the University of South Florida, St. Petersburg     

Organization of Plethodon salamander communities: guild-based community assembly

A long-standing goal in evolutionary ecology is to determine whether the organization of communities is reflective of underlying deterministic processes. In this study, I examined patterns of species co-occurrence among eastern Plethodon salamanders and determined whether they were consistent with predictions from a guild model of competition-based community assembly. Using a database of 45 species and 4540 geographic sites, I found that patterns of co-occurrence were significantly nonrandom at both a regional and continental scale, and species of different size guilds were distributed more evenly in sites than was expected by chance. Sites with the highest species richness had consistent patterns of community composition, and with few exceptions, the same five species were present at all sites. Taken together, these results imply that larger Plethodon communities are assembled from simpler communities in a manner consistent with what is predicted through competitive mechanisms and suggest that stable species combinations are possible to achieve at various levels of species richness. These results also provide strong evidence consistent with the hypothesis that competitive-based community assembly is a general phenomenon in Plethodon and that interspecific competition is prevalent among the eastern species of this group.     

A dynamic model of mating behavior in digger wasps: the energetics of male-male competition mimic size-dependent thermal constraints

I develop a state-based dynamic model of behavior to demonstrate that size-dependent differences in temperature tolerances are not necessary to account for the activity of small male digger wasps late in the day. In the model, males defend or patrol the nesting area, wait near nests, or feed away from the nesting area depending on time of day, energy reserves and size rank. I assume a large male competitive advantage, so mating opportunities decrease with size rank for territorial or patrolling males and are rare for all waiting males; the costs of patrolling or defense are higher than the costs of waiting. If energy reserves of all males are initially small, all males alternate feeding and territorial or patrolling behavior. If energy reserves are initially large, large males patrol or maintain territories until they risk starvation and leave the area to feed. At this time, smaller males that have conserved their resources by waiting and feeding may defend territories or patrol. I simulate the behavior of three populations representing two species of Microbembex by assuming large initial energy reserves for populations in which males were territorial and small initial reserves for populations in which males patrolled, and then convert the predicted time of activity to temperature using local regressions from field studies. Temporal patterns in the activity of large and small males were similar to those actually observed, and relationships between size and temperature predicted by the model corresponded to most observations and were sometimes positive. Thus, the delayed activity of smaller males does not correspond to activity at higher temperatures and is probably not attributable to size-dependent thermal tolerances, but may represent a temporal displacement of mating activity due to intra-sexual competition and mediated by energetics. The model makes testable predictions on the timing of feeding and depletion of energy reserves in relation to size and initial energy state, and suggests how differences among species may influence the temporal and spatial organization of male mating behavior. Received: 27 February 1997 / Accepted after revision: 26 July 1997     

The Reynolds transport theorem: Application to ecological compartment modeling and case study of ecosystem energetics

The Reynolds transport theorem (RTT) from mathematics and engineering has a rich history of success in mass transport dynamics and traditional thermodynamics. This paper introduces RTT as a complementary approach to traditional compartmental methods used in ecological modeling and network analysis. A universal system equation for a generic flow quantity is developed into a generic open-system differential expression for conservation of energy. Nonadiabatic systems are defined and incorporated into control volume (CV) and control surface (CS) perspectives of RTT where reductive assumptions in empirical data are then formally introduced, reviewed, and appropriately implemented. Compartment models are abstract, time-dependent systems of simultaneous differential equations describing storage and flow of conservative quantities between interconnected entities (the compartments). As such, they represent a set of flexible and somewhat informal, assumptions, definitions, algebraic manipulations, and graphical depictions subject to influence and selectively parsed expression by the modeler. In comparison, RTT compartment models are more rigorous and formal integro-differential equations and graphics initiated by the RTT universal system equation, forcing an ordered identification of simplifying assumptions, ending with clearly identified depictions of the transfer and transport of conservative substances in physical space and time. They are less abstract in the rigor of their equation development leaving less ambiguity to modeler discretion. They achieve greater consistency with other RTT compartment style models while possibly generating greater conformity with physical reality. Characteristics of the RTT approach are compared with those of a traditional compartment model of energy flow in an intertidal oyster-reef community.     

Influence of temperature and food availability on the ecological energetics of the giant scallop Placopecten magellanicus

Scallops (Placopecten magellanicus) were collected in 1981–1983 from two water depths at a location in Newfoundland, Canada, where temperature and food conditions associated with shallower water have been shown to be more favourable for somatic growth and gamete production. To gain insight into the seasonal energy balance for this species, metabolic and clearance rates were measured monthly under ambient temperature conditions and natural seston levels. Stereological techniques were used to determine the gamete volume fraction in the gonad in order to establish the annual reporductive cycle. The less favourable conditions associated with deeper water were relfected in reduced rates of gamete development, but the diameter of spawned eggs and the timing of spawning appeared unaffected by poorer conditions in the natural environment. Estimates of scope for growth were low or negative during the winter, but consistently high during the spring bloom, corresponding to a period of rapid gamete maturation. Somatic weight declined in both populations as gamete development proceeded but increased again during periods of low gametogenic activity, suggesting a close relationship between energy available for growth and the reproductive cycle. Oxygen uptake and clearance rate varied seasonally in relation to ambient temperature and food conditions, all of which appeared to be interrelated in a complex fashion with the energy demands of gametogenesis.     

Population structure and energetics of the shallow-water antarctic sea star Odontaster validus in contrasting habitats

Individuals and populations of Odontaster validus Koehler differed markedly among different habitats, as revealed in a study from October 1984 through January 1986 in McMurdo Sound, Antarctica. At McMurdo Station, individual sizes (wet weight) and population biomass (g wet wt m^-2 and kJ m^-2) decreased significantly with increasing depth. Individuals from shallow (10 to 20 m) habitats were in superior nutritional condition to those from deeper water (30 and 165 m), as shown by higher gonad and pyloric cecum indexes, and by higher lipid and energetic levels in the pyloric ceca. Moreover, gonadal output (reproductive output) was higher in shallow-water individuals. Higher levels of chlorophyll in the pyloric ceca and richer yellow to red coloration of the body wall in the shallow-water individuals indicate that they utilize the higher levels of primary production at shallow depths. At East Cape Armitage, where nearly permanent, thick, snow-covered ice most of the year resulted in very low levels of benthic primary production, the lowdensity sea stars were all very small and nutritionally similar to the deep-water individuals at McMurdo Station. At Cape Evans, where the generally snow-free sea-ice that broke up in mid-summer resulted in a luxurient benthic cover of diatoms and macroalgae, the sea stars were smaller than at McMurdo Station at comparable depths, but population densities were higher, resulting in 4 to 9 times greater biomass. Growth rates of sea stars fed in the laboratory were very low, especially compared to laboratory-reared temperate and tropical species; well-fed individuals need about 9 yr to reach 30 g wet weight, near the mean size of shallowwater individuals at McMurdo Station. No growth was detected in individuals caged at McMurdo Station for one year, suggesting even lower growth rates in the field. The stable size-frequency distributions at the different sites and depths throughout the year-long study suggest highly stable populations with low temporal variability in recruitment, migration and mortality. These data indicate that individuals and populations of O. validus quantitatively and qualitatively reflect the general level of productivity of a habitat. Differences noted in size, coloration, nutrition, and reproductive effort may be the result of long-term integration of local levels of primary production. These ubiquitous sea stars may serve as a biotic indicator of productivity in localized habitats around the continental shelf of Antarctica.     

Effect of suspended bottom material on growth and energetics in Mytilus edulis

The influence of suspended, natural silt (0 to 20 mg l^-1) in addition to unicellular algal cells (Phaeodactylum tricornutum) (o to 20.000 cells ml^-1) on clearance, growth and energetics in Mytilus edulis has been studied. Clearance increased by 32 to 43% by the addition of 5 mg silt l^-1 as compared to clearance in a pure algal suspension. Ingestion and growth rate increased with algal concentration, and growth rate was further increased by 30 to 70% by the addition of 5 mg silt l^-1. A growth rate comparable to maximum natural growth rates was reached only at the highest algal concentration in the presence of 5 mg siltl^-1. Assimilation efficiency of P. tricornutum decreased from 77% at 5,000 cells ml^-1 to 52% at 20,000 cells ml^-1. In the experiments with silt added, some 20 to 30% of the assimilated organic matter originated from the suspended bottom material. Net growth efficiency increased with growth rate at a decelerating rate, approaching a maximum of about 70%. It is concluded that suspended bottom material, which is always present in M. edulis' natural habitats, serves as an additional food source, and that M. edulis depends on suspended bottom material to exploit fully its clearance potential, and to reach the maximum growth rates observed in nature.     

Representing general theoretical concepts in structural equation models: the role of composite variables

Structural equation modeling (SEM) holds the promise of providing natural scientists the capacity to evaluate complex multivariate hypotheses about ecological systems. Building on its predecessors, path analysis and factor analysis, SEM allows for the incorporation of both observed and unobserved (latent) variables into theoretically-based probabilistic models. In this paper we discuss the interface between theory and data in SEM and the use of an additional variable type, the composite. In simple terms, composite variables specify the influences of collections of other variables and can be helpful in modeling heterogeneous concepts of the sort commonly of interest to ecologists. While long recognized as a potentially important element of SEM, composite variables have received very limited use, in part because of a lack of theoretical consideration, but also because of difficulties that arise in parameter estimation when using conventional solution procedures. In this paper we present a framework for discussing composites and demonstrate how the use of partially-reduced-form models can help to overcome some of the parameter estimation and evaluation problems associated with models containing composites. Diagnostic procedures for evaluating the most appropriate and effective use of composites are illustrated with an example from the ecological literature. It is argued that an ability to incorporate composite variables into structural equation models may be particularly valuable in the study of natural systems, where concepts are frequently multifaceted and the influence of suites of variables are often of interest.     

The effect of social interactions on calling energetics in the gray treefrog (Hyla versicolor)

Summary The vocal behavior of Hyla versicolor was studied in the field by means of behavioral observations and playback experiments, and these data were coupled with measurements of oxygen consumption in calling frogs to estimate the effect of social interactions on calling energetics. Male gray treefrogs have intense calls (median peak SPL=109 dB, fast RMS SPL=100 dB at 50 cm). At an air temperature of 23° C, males produced an average of 1,200–1,300 calls/h for 2–4 h per night. Calling rates and call durations differed among individuals, but were relatively constant for each male during periods of sustained calling. Males in dense choruses gave calls about twice as long as isolated males, but produced calls at about half the rate. Consequently, total calling effort and estimated aerobic costs were largely independent of chorus density. Playbacks of recorded calls to males in the field elicited increases in call duration and decreases in calling rate, regardless of the rate or duration of the stimulus. Males gave longer calls in response to long calls or to stimuli presented at high rates, but they did not precisely match either stimulus rate or duration. Calling effort and estimated oxygen consumption changed only slightly during stimulus playbacks. These results indicate that male-male competition elicits pro-found changes in the vocal behavior of calling males, but these changes have little effect on energy expenditure. We estimated that most calling males had metabolic rates of about 1.7–1.8 ml O₂/(g\h), or about 280 J/h for an average size (8.6 g) male at 20° C. Although changes in call duration and calling rate did not affect aerobic costs of calling, males producing long calls at slow rates called for fewer hours per night than males producing shorter calls at higher rates. This suggests that calling time may be limited by the rate at which muscle glycogen reserves are depleted.     

Influence of the water-accommodated fraction of no. 2 fuel oil on energetics of Cancer irroratus larvae

The influence of the water-accommodated fraction of No. 2 fuel oil on aspects of larval development of Cancer irroratus (Say) has been investigated. Rates of survival, respiration, ingestion and growth were estimated for both oil-treated and untreated larvae. Larvae reared in oil-treated water exhibited lower survival to megalopa than did larvae reared in untreated water. The presence of oil also altered several aspects of larval energetics. Reductions were seen in food consumption and growth rate while the metabolic cost of maintenance increased. One overall effect that may occur as a result of lowered larval survival and growth rate would be the reduction in larval recruitment in oil-polluted areas.     

Reproductive energetics of two species of dorid nudibranchs with planktotrophic and lecithotrophic larval strategies

Onchidoris muricata (Müller) and Adalaria proxima (Alder and Hancock) are sympatric, potentially competing species of dorid nudibranchs, which preferentially graze the cheilostome polyzoan Electra pilosa (L.). O. muricata is small and lays small eggs which hatch as poorly-developed planktotrophic veliger larvae. A. proxima is larger and reproduces by means of larger eggs which hatch, as well-developed lecithotrophic larvae, that can metamorphose within approximately 24 h of release. A. proxima larvae can feed in the plankton, but do not require extrinsic nutrition to undergo complete development. Both species spawn in February–april, and have a strictly annual life-cycle. Comparisons of the calorific content of spawn have shown that A. proxima apportions a greater number of calories to reproduction, but that O. muricata makes a greater relative effort. A. proxima shown considerable individual variability in reproductive effort, which fails to correlate with, body size or rate of spawning. A more deterministic situation applies to O. muricata, because body size and fecundity follow an allometric relationship. It appears that there is a threshold of absolute energy required to support the lecithotrophic larval strategy in nudibranchs, and that this is not attained by the smaller species, O. muricata. A. proxima thus appears to have both reproductive strategies open to it, and to have adopted lecithotrophy in order to offset the unpredictability of energy available for reproduction.Communicated by J. M. Mauchline, Oban     

A model for energetics and bioaccumulation in marine mammals with applications to the right whale.

We present a dynamic energy budget (DEB) model for marine mammals, coupled with a pharmacokinetic model of a lipophilic persistent toxicant. Inputs to the model are energy availability and lipid-normalized toxicant concentration in the environment. The model predicts individual growth, reproduction, bioaccumulation, and transfer of energy and toxicant from mothers to their young. We estimated all model parameters for the right whale; with these parameters, reduction in energy availability increases the age at first parturition, increases intervals between reproductive events, reduces the organisms' ability to buffer seasonal fluctuations, and increases its susceptibility to temporal shifts in the seasonal peak of energy availability. Reduction in energy intake increases bioaccumulation and the amount of toxicant transferred from mother to each offspring. With high energy availability, the toxicant load of offspring decreases with birth order. Contrary to expectations, this ordering may be reversed with lower energy availability. Although demonstrated with parameters for the right whale, these relationships between energy intake and energetics and pharmacokinetics of organisms are likely to be much more general. Results specific to right whales include energy assimilation estimates for the North Atlantic and southern right whale, influences of history of energy availability on reproduction, and a relationship between ages at first parturition and calving intervals. Our model provides a platform for further analyses of both individual and population responses of marine mammals to pollution, and to changes in energy availability, including those likely to arise through climate change.     

Disturbance and resource availability act differently on the same suite of plant traits: revisiting assembly hypotheses

Understanding the mechanisms of trait selection at the scale of plant communities is a crucial step toward predicting community assembly. Although it is commonly assumed that disturbance and resource availability constrain separate suites of traits, representing the regenerative and established phases, respectively, a quantification and test of this accepted hypothesis is still lacking due to limitations of traditional statistical techniques. In this paper we quantify, using structural equation modeling (SEM), the relative contributions of disturbance and resource availability to the selection of suites of traits at the community scale. Our model specifies and reflects previously obtained ecological insights, taking disturbance and nutrient availability as central drivers affecting leaf, allometric, seed, and phenology traits in 156 (semi-) natural plant communities throughout The Netherlands. The common hypothesis positing that disturbance and resource availability each affect a set of mutually independent traits was not consistent with the data. Instead, our final model shows that most traits are strongly affected by both drivers. In addition, trait-trait constraints are more important in community assembly than environmental drivers in half of the cases. Both aspects of trait selection are crucial for correctly predicting ecosystem processes and community assembly, and they provide new insights into hitherto underappreciated ecological interactions.     

On the intrinsic structure of differential equation models of ecosystems

Some intrinsic properties of differential equation models of ecosystems are formulated. The properties, which are long-termed, are classified into fundamental, stability, and sensitivity substructures. An aspect of the ecological term “resilient” — asymptotic stability uniformly for all small parameter variations — is introduced in a mathematical setting. In the sensitivity substructure, bounded sensitivity and continuity of sensitivity with respect to solutions are recognized as intrinsic properties. Illustrations of these two properties are given.     

Feeding ecology and energetics of the Antarctic chaetognaths Eukrohnia hamata, E. bathypelagica and E. bathyantarctica

Ecological and physiological studies focused on dietary preferences, lipid biochemistry and energetics within the three Antarctic chaetognaths Eukrohnia hamata, E. bathypelagica and E. bathyantarctica from meso- and bathypelagic depths. Eukrohnia hamata and E. bathypelagica respired 0.15 μL O₂ mg dry mass (DM)⁻¹ h⁻¹, which translates to an average metabolic loss of only <1.1% of body carbon per day. Lipid storage was not substantial in E. bathypelagica (mean 11.5 ± 6.5% DM) and E. bathyantarctica (mean 15.4 ± 4.1% DM) during summer and winter, suggesting year-round feeding of these predators mainly on copepods. In E. bathypelagica, total fatty acids were dominated by the fatty acids 16:0, 20:5(n-3) and 22:6(n-3) and in E. bathyantarctica also by 18:1(n-9), a fatty acid usually found in storage lipids. Only the latter species was characterized by significant amounts of wax esters, consisting largely of the common fatty alcohols 16:0, 20:1(n-9) and the unusual fatty alcohol isomer 22:1(n-9).