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.     

Biokinetics of plutonium,americium and californium in the marine isopod Cirolana borealis,with observations on its feeding and molting behavior

In a series of laboratory experiments carried out during 1982, the biokinetics of ²³⁷Pu (IV), ²³⁷Pu (V), ²⁴¹Am (III), and ²⁵²Cf (III) were studied in the benthic marine isopod Cirolana borealis Lilljeb. Following a 3 wk exposure to labelled seawater, isopods reached non-equilibrium concentration factors of 52±15, 54±12, 176±14, and 185±53 for Pu (IV), Pu (V), Am and Cf, respectively. Uptake of these radionuclides occurred to a great extent by surface adsorption, with the major fraction of the accumulated radionuclides being associated with exoskeleton and lesser percentages located in gut, digestive gland, muscle and haemolymph. The pattern of radiotracer depuration was in all cases biphasic, with the long-lived retention compartment retaining the largest fraction of whole-body radioactivity. Biological half-lives for radionuclide turnover in the long-lived compartment were 60±3 d, 87±5 d, 261±12 d and 288±60 d for Pu (IV), Pu (V), Am and Cf, respectively. Examination of radionuclide distribution among tissues following longterm depuration showed that the radionuclides were eliminated from internal tissues more rapidly than from the exoskeleton. Uptake and depuration studies indicated that the biokinetics of americium and californium were similar to each other, but that they differed significantly from those of oxidized and reduced plutonium. In general, these experimental results support previous reports that americium and californium are more bioavailable than plutonium in the marine waters. The fraction americium and californium assimilated into tissue from food was very low and did not exceed 5%. Unassimilated labelled food was often retained in the gut for more than 2 mo and released only infrequently in a loosely bound condition. In the event of contamination, this particular feeding-digestion strategy in highly mobile, scavenging isopods could act as a potential biological mechanism for the widespread dispersion of radioactive wastes.     

A finite element method to solve a population dynamics stage-structured model of intertidal barnacles

Many marine organisms are fixed or highly sedentary as adults but the adult population may be strongly dependent on the oceanic transport of planktonic larvae. In order to assess interactions between oceanographic and biological processes that determine the population dynamics of marine organisms with a sessile adult phase restricted to the coastline and a planktonic larval phase, we present a stage-structured finite element model for the barnacle Balanus glandula that inhabits the rocky intertidal zone of central California, USA.     

Human trampling as short-term disturbance on intertidal mudflats: effects on macrofauna biodiversity and population dynamics of bivalves

The effect of physical disturbance in the form of trampling on the benthic environment of an intertidal mudflat was investigated. Intense trampling was created as unintended side-effect by benthic ecologists during field experiments in spring and summer 2005, when a mid-shore area of 25 × 25 m was visited twice per month by on average five researchers for a period of 8 months. At the putatively-impacted location (I) (25 × 25 m) and two nearby control locations (Cs) (25 × 25 m each), three sites (4 × 4 m) were randomly selected and at each site, three plots (50 × 50 cm) were sampled after 18 and 40 days from the end of the disturbance. Multivariate and univariate asymmetrical analyses tested for changes in the macrofaunal assemblage, biomass of microphytobenthos and various sediment properties (grain-size, water content, NH₄ and NO₃ concentrations in the pore water) between the two control locations (Cs) and the putatively-impacted location (I). There were no detectable changes in the sediment properties and microphytobenthos biomass, but variability at small scale was observed. Microphytobenthos and NH₄ were correlated at I to the number of footprints, as estimated by the percentage cover of physical depressions. This indicated that trampling could have an impact at small scales, but more investigation is needed. Trampling, instead, clearly modified the abundance and population dynamics of the clam Macoma balthica (L.) and the cockle Cerastoderma edule (L.). There was a negative impact on adults of both species, probably because footsteps directly killed or buried the animals, provoking asphyxia. Conversely, trampling indirectly enhanced recruitment rate of M. balthica, while small-sized C. edule did not react to the trampling. It was likely that small animals could recover more quickly because trampling occurred during the growing season and there was a continuous supply of larvae and juveniles. In addition, trampling might have weakened negative adult-juvenile interactions between adult cockles and juvenile M. balthica, thus facilitating the recruitment. Our findings indicated that human trampling is a relevant source of disturbance for the conservation and management of mudflats. During the growing season recovery can be fast, but in the long-term it might lead towards the dominance of M. balthica to the cost of C. edule, thereby affecting ecosystem functioning.     

Vertical swimming in wave-induced currents as a control mechanism of intertidal migration by a sand-beach isopod

Both the vertical and horizontal distributions ofEurydice pulchra (Leach) within the water column of the surf zone were recorded throughout complete tidal cycles on a sandy beach in North Wales during the summer of 1989. Upon emerging from the sand, the isopods tended to swim up in the water column, where transport onshore would be facilitated by the wave-induced, onshore currents which laboratory wave-tank experiments have confirmed to occur near the water surface. This combination of active and passive transport to the water's edge results in high numbers of individuals in the narrow swash zone. At and just after the time of high tide, individuals swim to the water/sediment interface where, as again confirmed by wave-tank experiments, the predominant water movement is offshore. Continued swimming near the bottom during the ebb tide before reburrowing in the sand ensures transport downshore and avoidance of stranding above the characteristic level of zonation on the shore. Vertical migrations ofE. pulchra in the water column permit differential exploitation of up and downshore currents to achieve horizontal migration.     

Infestation dynamics of gnathiid isopod juveniles parasitic on the coral-reef fish Hemigymnus melapterus (Labridae)

The dynamics of infestation by parasitic juveniles of gnathiid isopods were investigated at Lizard Island in the summer of 1997/1998 to determine when, and at what rate, they infest fishes. Variation in gnathiid abundance on wild-caught fish (Hemigymnus melapterus) between dawn and sunset was examined, and unparasitized H. melapterus in cages were exposed to gnathiids in the field for 4 h (8 h for fish sampled at 06:00 h) at five different times of the day and night (10:00, 14:00, 18:00, 22:00, 06:00 hrs) on three reefs. To control for any potential effect of the cleaner-fish Labroides dimidiatus, which consumes large numbers of gnathiids each day, sampling was carried out on reefs from which all cleaner-fish had been removed. Gnathiid abundance decreased during the day. Standardized abundance per wild-caught fish was 1.9 times higher at dawn than at sunset. Gnathiids successfully infested fish in cages. Sixty-one percent of the fish in cages were infested with ≥1 gnathiids, with 51% of the fish having between 1 and 4 gnathiids after 4 h exposure. Gnathiids infested fish both day and night. The log₁₀ (abundance +1) of gnathiids per caged fish varied significantly between time periods, with higher gnathiid numbers on fish sampled at 18:00, 22:00, and 06:00 hrs than on fish sampled at 10:00 and 14:00 hrs. In contrast to gnathiid abundance on wild-caught fish, the number of gnathiids on caged fish sampled at sunset was as high as that at dawn. The estimated mean (SE) cumulative number of gnathiids per caged fish [mean size = 13.7 cm (±0.25)] per day was 7.8 (1.1); this is similar to the estimated mean number of 7.3 gnathiids on similar-sized wild-caught fish at 6:00 hrs. The high infestation rate of gnathiids on caged fish in the late afternoon contrasted with the low numbers on wild-caught fish at this time, suggesting that factors other than infestation behaviour may be responsible for the low numbers of gnathiids on wild-caught fish in the afternoon. Received: 19 August 1998 / Accepted: 1 June 1999     

Population dynamics and reproductive biology of the commensal isopod Colidotea rostrata (Crustacea: Isopoda: Idoteidae)

Colidotea rostrata (Benedict, 1898) is the only known commensal idoteid isopod, living on and mimicking the color of two northeastern Pacific sea urchins of the genus Strongylocentrotus. The population dynamics and reproductive biology of C. rostrata on its host urchins were studied at a low rocky intertidal area in southern California (33°40N; 118°30W) from December 1984 to December 1986. Isopod populations remained relatively stable throughout the 2 yr study, with isopods inhabiting an average of 56.1% of the urchins at 6.1±0.6 (x±2 SE) isopods per urchin. Female isopods reach sexual maturity at a length of 6.8 mm. Fecundity in C. rostrata averaged 11.8±0.9 (x±2 SE) embryos per brood, and increased with female body size. Breeding occurs all year long in C. rostrata, with a main reproductive period between the warmer spring and summer months. Newly released mancas and juvenile isopods were present during all months of the study. C. rostrata differs from the free-living Idoteidae in its smaller maximum size, reduced fecundity, 1:1 sex ratio, and low juvenile mortality. These features may represent adaptations to a commensal life style that reflect a reduced mortality pressure on these isopods.     

Interacting trophic forcing and the population dynamics of herring

Small pelagic fish occupy a central position in marine ecosystems worldwide, largely by determining the energy transfer from lower trophic levels to predators at the top of the food web, including humans. Population dynamics of small pelagic fish may therefore be regulated neither strictly bottom-up nor top-down, but rather through multiple external and internal drivers. While in many studies single drivers have been identified, potential synergies of multiple factors, as well as their relative importance in regulating population dynamics of small pelagic fish, is a largely unresolved issue. Using a statistical, age-structured modeling approach, we demonstrate the relative importance and influence of bottom-up (e.g., climate, zooplankton availability) and top-down (i.e., fishing and predation) factors on the population dynamics of Bothnian Sea herring (Clupea harengus) throughout its life cycle. Our results indicate significant bottom-up effects of zooplankton and interspecific competition from sprat (Sprattus sprattus), particularly on younger age classes of herring. Although top-down forcing through fishing and predation by grey seals (Halichoerus grypus) and Atlantic cod (Gadus morhua) also was evident, these factors were less important than resource availability and interspecific competition. Understanding key ecological processes and interactions is fundamental to ecosystem-based management practices necessary to promote sustainable exploitation of small pelagic fish.     

Ungulate population dynamics and optimization models

Optimal harvesting strategies for an ungulate population are estimated using stochastic dynamic programming. Data on the Llano Basin white-tailed deer (Odocoileus virginianus) population were used to construct a 2-variable population dynamics model. The model provided the basis for estimating optimal harvesting strategies as a feedback function of the current values of the state variables (prefawning older deer and juveniles). Optimal harvest strategies were insensitive to assumptions about the probability distributions of the stochastic variable (rainfall). The response of the population components to harvesting and the returns obtained from applying optimal strategies were explored through simulation. Mean annual harvest is about 15% of the population. Simplified harvesting strategies based on age-ratios as well as a simplified version based on optimal strategies—but assuming persisting equilibrium juvenile deer density—were compared to optimal strategies through examining values of information. Simplified harvesting strategies lead to a lower harvest over a 50-year simulation period.     

Hydrology influences population genetic structure and connectivity of the intertidal amphipod Corophium volutator in the northwest Atlantic

The mechanisms driving genetic structure in marine systems are elusive due to the difficulty of identifying temporal and spatial barriers to dispersal. By studying marine invertebrate species with limited dispersal potential, genetic structure can be directly related to physical and biological factors restricting connectivity. In the northwest Atlantic, the benthic brood-rearing amphipod Corophium volutator is distributed across basins with distinct circulation patterns and has the potential to disperse passively during its adult stage. We analyzed spatial genetic variation and migration rates across C. volutator’s North American range with sequence data for mitochondrial DNA and three novel nuclear markers using frequency and coalescent-based methods. We found low genetic differentiation within basins, but strong subdivision within the Bay of Fundy and a striking biogeographic break between the Bay of Fundy and Gulf of Maine, suggesting that genetic drift may act on populations in which connectivity is restricted due to the limitation of passive dispersal by hydrological patterns.     

Growth dynamics and mortality of the intertidal encrusting sponge Halichondria okadai (Demospongiae, Halichondrida)

A periodical survey of the growth of an intertidal encrusting sponge Halichondria okadai was conducted on a rocky intertidal shore on the Izu Peninsula, southern Japan. The area covered by each of 16 marked sponges was monitored, and the fusion and fission of specimens were recorded from June 1995 to June 1998. During the investigation, 15 of the original specimens managed to survive accompanying fusions or fissions and only one colony completely disappeared. Ten fusions and 23 fissions were observed in total. The maximum death rate during a semilunar period was 16.7% and cumulative mortality throughout the study period was 46.1%, indicating that the mortality of H. okadai is lower than that of other species previously reported. There was no particular seasonal pattern in the occurrences of fusion, fission and death of sponges. However, fissions were frequently observed in the latter half of the study period and were sometimes followed by fusions or death of specimens. Positive growth rates were often observed during warmer months, and specimens showed little growth or regression in winter. Seasonal patterns in surface area were thought to be related to seasonality in reproductive activity.     

Energy flow and population dynamics of the barnacle Balanus glandula

The energetics and population dynamics of a barnacle (Balanus glandula Darwin) population in British Columbia, Canada, were studied. Consumption, energy flow, production and mortality were 6844.6, 6667.0, 2896.5 and 2522.8 Kcal m^-2 year^-1, respectively. These energy flow and production values are among the highest for animal populations reported, and therefore strongly suggest the functional importance of E. glandula in littoral systems. The young age groups of the first-year settlements were most important in contributing to the energy flow, production and reproduction of the entire population. Most of the assimilated energy in the older age groups was used in respiration.     

Changes in barnacle population structure along an intertidal community gradient in the Florida Keys

Population distributions, densities and percentage deaths and individual size variations of the barnacles Chthamalus stellatus and Tetraclita squamosa, and the distribution of the vermetid snail Spirogluphus annulatus are measured to clarify determinants the Florida Keys (USA). The species composition of the pilings community and barnacle population densities correspond to physical gradients dependent on piling distance from shore. Interspecific competition for space is insignificant in determining C. stellatus presence or absence on pilings. Intraspecific crowding in highdensity zones of adult T. squamosa is evidenced by a significant decrease in basal diameter as density increases. The peak percentage of dead C. stellatus coincides with peak densities. The percentage of dead T. squamosa is at a low point at peak densities due to the ability of T. squamosa cyprids to settle and survive on adult shells in high-density regions. Feasible perturbation experiments for testing the importance of interspecific competition in determining the densities and vertical distributions of the species are discussed. The results of such experiments can be used to test the assumptions and predictions of the widely applied competition coefficient measure proposed by Levins (1968). Several theoretical deficiencies of Levins' measure are discussed.     

Uncertainty analysis of transient population dynamics

Two types of demographic analyses, perturbation analysis and uncertainty analysis, can be conducted to gain insights about matrix population models and guide population management. Perturbation analysis studies how the perturbation of demographic parameters (survival, growth, and reproduction parameters) may affect the population projection, while uncertainty analysis evaluates how much uncertainty there is in population dynamic predictions and where the uncertainty comes from. Previously, both perturbation analysis and uncertainty analysis were conducted on the long-term population growth rate. However, the population may not reach its equilibrium state, especially when there is management by harvesting or hunting. Recently, there has been an increased interest in short-term transient dynamics, which can differ from asymptotic long-term dynamics. There are currently techniques to conduct perturbation analyses of short-term transient dynamics, but no techniques have been proposed for uncertainty analysis of such dynamics. In this study, we introduced an uncertainty analysis technique, the general Fourier Amplitude Sensitivity Test (FAST), to study uncertainties in transient population dynamics. The general FAST is able to identify the amount of uncertainty in transient dynamics and contributions by different demographic parameters. We applied the general FAST to a mountain goat (Oreamnos americanus) matrix population model to give a clear illustration of how uncertainty analysis can be conducted for transient dynamics arising from matrix population models.     

Vertical zonation of endosymbiotic zooxanthellae within a population of the intertidal sea anemone,Anthopleura uchidai

Intertidal organisms commonly form zonation bands along the shore. Environmental stressors often determine the vertical position of each zonation band. These stressors may similarly affect the distribution pattern of endogenous species in their intertidal hosts. To evaluate this possibility, we investigated the distribution pattern of endosymbiotic zooxanthellae in the genus Symbiodinium in a population of the intertidal sea anemone Anthopleura uchidai. We used molecular genetics to identify the Symbiodinium clades and found that A. uchidai has two clades of Symbiodinium, clades A and F. These Symbiodinium clades were disproportionally distributed along the vertical gradient of the intertidal shore. Anemones on the upper shore exclusively possessed clade F Symbiodinium while clade A Symbiodinium became dominant in the sea anemones on the lower shore. Photosynthesis activity assays showed that these Symbiodinium clades had similar net productivities at 23.3 and 31.8 °C at all irradiance levels. At 35 °C, however, clade A Symbiodinium exhibited substantially lower net productivities than clade F Symbiodinium, demonstrating that these Symbiodinium clades have distinct tolerances to thermal stress. These results suggest that the thermal gradient across tidal height is a major factor shaping the zonation pattern of Symbiodinium clades in A. uchidai.     

Analysis of an intertidal population of the amphipod Gammarus palustris using a modified version of the egg-ratio method

An intertidal population of Gammarus palustris Bousfield was sampled over a 23-month period and analyzed by a modified form of the egg-ratio method. Total numbers and size-frequency distributions revealed small numbers in the early spring which later increased to a peak in the summer months. This was followed by a marked reduction near the end of sumer and a subsequent rise in numbers during the early fall. Finally there was a second decrease to very low numbers in the late fall and early winter. Average brood size was calculated and used as an indication of food limitation. Attempts to correlate this with per capita rate of change (r), birth rate ( \((\hat b)\) and death rate ( \((\hat d)\) ) suggested that the population increase in the early summer was food-rather than predator-limited. The late summer decline appears to be the result of predation pressure as well as food limitation. The fall peak is attributed to juvenile release from ovigerous femals escaping previous predation and to an increase in adult males resulting from a release in predation pressure. The winter decline may be due to adverse environmental conditions.     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type.     

Comparisons of genetic population structures in four intertidal brachyuran species of contrasting habitat characteristics

Genetic population structures along the Japanese coast, analyzed by sequence data from the mitochondrial DNA COI region, were determined for four intertidal brachyuran species in the superfamily Thoracotremata (Ocypode ceratophthalma, Gaetice depressus, Chiromantes dehaani and Deiratonotus japonicus), which were characterized by different habitat requirements. O. ceratophthalma (seashore; supratidal sand) and C. dehaani (estuarine; supratidal marsh) showed no significant genetic differentiation among Japanese populations. The Japanese populations of O. ceratophthalma, however, were found to genetically differentiated from the Philippine population. G. depressus (seashore; intertidal cobbles) exhibited significant genetic differentiation between the Amami-Ohshima population and other local populations. D. japonicus (estuarine; intertidal cobbles) showed significant genetic differentiation among many local populations separated by about 30–1,200 km. The different patterns of genetic population structure recorded for the four species, thus, do not simply correspond to habitat type. An erratum to this article can be found at