Precopulatory guarding-time of the male amphipod Eogammarus oclairi: effect of population structure

This study shows that population structure affects guarding time in the amphipod Eogammarus oclairi Bousfield. In the laboratory, mean guarding time depended on the male weight-frequency distribution, operational sex ratio, and male density. It was longer when the male population encompassed a broad range of weights than when the weight distribution was narrower; however, guarding time did not differ when the population's weight distribution was narrow and composed of either small or large males. Mean guarding time varied between a few hours when the operational sex ratio (male:female) was 1:3 to>7 d when the sex ratio was 2:1. It was also affected by male density, increasing as male density increased. The results suggest that the stimulus to guard depends on the intensity of male-male competition.     

Habitat selection in the tropical polychaeteSpirobranchus giganteus

Total body weight, tube length, abdomen weight and branchial crown weight of individualSpirobranchus giganteus (Pallas) living on four different coral species on the bank reef on the west coast of Barbados were investigated in 1986. Worms onDiploria strigosa were larger in all size parameters than those onMontastrea annularis, which were larger than those onMontastrea cavernosa, which were larger than those onPorites porites. The differences between worms onM. annularis andM. cavernosa were not significant. Variation in worm size on the different corals could result from variation in mortality and/or variation in growth. Whichever the case, larger worms may have higher lifetime reproductive success. Abdomen weight is an index of gamete production inS. giganteus, and increases with increasing body weight for worms on all corals. Moreover, larger worms do not have lower abdomen weight at a given body size than smaller worms. This may imply that faster growth does not reduce gamete production at body size. The coral species on which worms are largest are those most preferred by worm larvae in the laboratory and most heavily colonised by adult worms in the field. The results suggest adaptive habitat selection by planktonic larvae ofS. giganteus.     

Reproductive success,relative abundance and population structure of two species of Nephtys in an estuarine beach

During 1978 and 1979 specimens of Nephtys hombergii in the River Tyne estuary (North East England) became gravid during the winter but did not spawn in the spring breeding season; instead their gametes were resorbed. The drastic effects of these spawning failures were apparent in the population structure in 1980, when the 1978 and 1979 year-classes were shown to be virtually absent, by the size of the individuals in the population and the numbers of growth lines in their jaws. The population structure in 1975, by contrast, indicated that recruitment during the preceding 4 to 5 yr had been good. Spawning was also effective in 1980, and an 0 group was detected in August 1980. Comparable data has been obtained for N. caeca, which occurs in the same beach. It shows that recruitment of this species during 1976–1980 was consistently good, whereas analysis of the age-class frequencies suggests that recruitment of N. caeca was poor in 1973 and 1975. The dominance relationship of the two species in the beach has been reversed during the period 1975–1980. It is suggested that the sympatric distribution of the two species is maintained in part by periodic reproductive failure. The causes of this are not yet known; possible reasons, including the extremely cold conditions experienced in 1978–1979 are discussed. The difference in frequency of the 1978, 1979 and 1980 year classes will now permit a rigorous evaluation of the relationship between the number of growth lines in the jaws and age in N. hombergii.     

Polygyny and extra-pair paternity enhance the opportunity for sexual selection in blue tits

Polygyny and extra-pair paternity are generally thought to enhance sexual selection. However, the extent to which these phenomena increase variance in male reproductive success will depend on the covariance between success at these two strategies. We analysed these patterns over four breeding seasons in facultatively polygynous blue tits Cyanistes caeruleus. We found that both polygyny and extra-pair paternity increased variance in male reproductive success and that standardised variance in annual number of genetic fledglings was 2.6 times higher than standardised variance in apparent success when assuming strict monogamy. Nevertheless, male success at securing within-pair paternity was unrelated to success at gaining extra-pair paternity and, when considering the positive effect of age on extra-pair success and attracting a second female, polygynous males were no more likely to sire extra-pair fledglings. Overall, polygynous males fledged more genetic offspring than monogamous males, but first-year polygynous males lost a greater share of within-pair paternity. A literature review suggests that this adverse effect of polygyny on within-pair paternity is frequent among birds, inconsistent with the prediction that females engage in extra-pair copulation with successful males to obtain good genes. Furthermore, a male's share of paternity was repeatable between years, and among females of polygynous males within years, such that a compatibility function of extra-pair copulations was likewise unsupported. Instead, we suggest that the observed patterns are most consistent with a fertility insurance role for extra-pair copulations, which does not exclude the greater opportunity for sexual selection through differential ability of males to gain paternity.     

Effects of predation and habitat structure on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Caridea) on temperate rocky reefs

Human disturbances, such as overfishing, may disrupt predator–prey interactions and modify food webs. Underwater surveys were carried out at six shallow-water reef barrens in temperate waters of northern-central Chile from October to December 2010 to describe the effects of predation, habitat complexity (low, medium and high) and refuge availability on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Rhynchocinetidae), an important mesoconsumer on subtidal hard substrata. Three sites were within managed (restricted access) areas for fishermen, and three were unmanaged (open-access). Field observations and tethering experiments were conducted to examine the relationship between fish and shrimp abundances, and the relative predation rates on shrimps. Direct effects of predation on R. typus body-size distribution were examined from shrimps collected in the field and fish stomachs. The presence and the abundance of R. typus increased with habitat reef complexity and refuge availability. Shrimp abundance was negatively related to fish abundance in managed areas, but not in open-access areas, where shrimp densities were the highest. Also, predation rates and body-size distribution of shrimps were unrelated, although fish consumed more large shrimps than should be expected from their distribution in the field. R. typus occurred most often in shelters with wide openings, offering limited protection against predators, but providing potential aggregation sites for shrimps. Overall, direct effects of predation on shrimp densities and population structure were weak, but indirect effects on shrimp distribution within reefs appear to have been mediated through behavioural responses. Our study highlights the need to assess both numerical and behavioural responses of prey to determine the effects of predator loss on mesoconsumer populations.     

Spatial scaling of avian population dynamics: population abundance, growth rate, and variability

Synchrony in population fluctuations has been identified as an important component of population dynamics. In a previous study, we determined that local-scale (<15-km) spatial synchrony of bird populations in New England was correlated with synchronous fluctuations in lepidopteran larvae abundance and with the North Atlantic Oscillation. Here we address five questions that extend the scope of our earlier study using North American Breeding Bird Survey data. First, do bird populations in eastern North America exhibit spatial synchrony in abundances at scales beyond those we have documented previously? Second, does spatial synchrony depend on what population metric is analyzed (e.g., abundance, growth rate, or variability)? Third, is there geographic concordance in where species exhibit synchrony? Fourth, for those species that exhibit significant geographic concordance, are there landscape and habitat variables that contribute to the observed patterns? Fifth, is spatial synchrony affected by a species' life history traits? Significant spatial synchrony was common and its magnitude was dependent on the population metric analyzed. Twenty-four of 29 species examined exhibited significant synchrony in population abundance: mean local autocorrelation (rho)= 0.15; mean spatial extent (mean distance where rho=0) = 420.7 km. Five of the 29 species exhibited significant synchrony in annual population growth rate (mean local autocorrelation = 0.06, mean distance = 457.8 km). Ten of the 29 species exhibited significant synchrony in population abundance variability (mean local autocorrelation = 0.49, mean distance = 413.8 km). Analyses of landscape structure indicated that habitat variables were infrequent contributors to spatial synchrony. Likewise, we detected no effects of life history traits on synchrony in population abundance or growth rate. However, short-distance migrants exhibited more spatially extensive synchrony in population variability than either year-round residents or long-distance migrants. The dissimilarity of the spatial extent of synchrony across species suggests that most populations are not regulated at similar spatial scales. The spatial scale of the population synchrony patterns we describe is likely larger than the actual scale of population regulation, and in turn, the scale of population regulation is undoubtedly larger than the scale of individual ecological requirements.     

Mesozooplankton community structure, abundance and biomass in the central Arctic Ocean

During the "International Arctic Ocean Expedition 1991" (20 August-21 September 1991) mesozooplankton was sampled at six stations in the Nansen, Amundsen and Makarov Basins of the central Arctic Ocean from 1,500 m depth to the surface by multiple opening/closing net hauls. Total mesozooplankton abundance decreased from 268 ind. m^-3 in the surface layer (0-50 m) to <25 ind. m^-3 below 200 m depth. The small copepods Oithona similis and Microcalanus pygmaeus, as well as copepod nauplii, were most abundant close to the surface, while Oncaea borealis and Spinocalanus spp. frequently occurred at greater depth. Mesozooplankton dry mass (DM) integrated over the upper 1,500 m of the water column was surprisingly stable throughout the investigation area and measured 2.0ǂ.3 g DM m^-2. Dry mass in the upper 50 m measured 20.9 mg m^-3 and was dominated by Calanus hyperboreus (57.4%) and C. glacialis (21.1%). C. finmarchicus was very abundant only in the Nansen Basin. Below 200 m the calanoid copepods Metridia longa, Microcalanus pygmaeus and Pareuchaeta spp., the decapod Hymenodora glacialis and chaetognaths of the genus Eukrohnia were the principal contributors to biomass values of <1 mg DM m^-3. Hence, vertical changes in abundance, biomass and species composition were much more pronounced than regional differences between the basins. Three different mesozooplankton communities were differentiated according to their faunistic composition and are discussed in context with the major water masses: Polar Surface Water, Atlantic Layer and Arctic Deep Water.     

Demographic heterogeneity, cohort selection, and population growth

Demographic heterogeneity--variation among individuals in survival and reproduction--is ubiquitous in natural populations. Structured population models address heterogeneity due to age, size, or major developmental stages. However, other important sources of demographic heterogeneity, such as genetic variation, spatial heterogeneity in the environment, maternal effects, and differential exposure to stressors, are often not easily measured and hence are modeled as stochasticity. Recent research has elucidated the role of demographic heterogeneity in changing the magnitude of demographic stochasticity in small populations. Here we demonstrate a previously unrecognized effect: heterogeneous survival in long-lived species can increase the long-term growth rate in populations of any size. We illustrate this result using simple models in which each individual's annual survival rate is independent of age but survival may differ among individuals within a cohort. Similar models, but with nonoverlapping generations, have been extensively studied by demographers, who showed that, because the more "frail" individuals are more likely to die at a young age, the average survival rate of the cohort increases with age. Within ecology and evolution, this phenomenon of "cohort selection" is increasingly appreciated as a confounding factor in studies of senescence. We show that, when placed in a population model with overlapping generations, this heterogeneity also causes the asymptotic population growth rate lambda to increase, relative to a homogeneous population with the same mean survival rate at birth. The increase occurs because, even integrating over all the cohorts in the population, the population becomes increasingly dominated by the more robust individuals. The growth rate increases monotonically with the variance in survival rates, and the effect can be substantial, easily doubling the growth rate of slow-growing populations. Correlations between parent and offspring phenotype change the magnitude of the increase in lambda, but the increase occurs even for negative parent-offspring correlations. The effect of heterogeneity in reproductive rate on lambda is quite different: growth rate increases with reproductive heterogeneity for positive parent-offspring correlation but decreases for negative parent-offspring correlation. These effects of demographic heterogeneity on lambda have important implications for population dynamics, population viability analysis, and evolution.     

Habitat selection models to account for seasonal persistence in radio telemetry data

Models for the analysis of habitat selection data incorporate covariates in an independent multinomial selections model (McCracken et al. 1998) Ramsey and Usner 2003 and an extension of that model to include a persistence parameter (2003). In both cases, all parameters are assumed to be fixed through time. Radio telemetry data collected for habitat selection studies typically consist of animal relocations through time, suggesting the need for an extension to these models. We use a Bayesian approach that allows for the habitat selection probabilities, persistence parameter, or both, to change with season. These extensions are particularly important when movement patterns are expected to differ seasonally and/or when availabilities of habitats change throughout the study period due to weather or migration. We implement and compare the models using radio telemetry data for westslope cutthroat trout in two streams in eastern Oregon.     

Changes in body size, abundance, and energy allocation in rockfish assemblages of the northeast Pacific.

Fish body size, a key driver of many aspects of fish population biology and ecology, is affected by fisheries that deplete the largest individuals. Rockfish (genus Sebastes) are a diverse group that has been heavily fished on the U.S. West Coast in recent decades. We examined trawl survey data from 1980 to 2001 to determine spatial and temporal trends in body size and density of 16 shelf rockfish species, including six that are considered overfished. Mean individual mass and maximum observed mass declined in the majority of species in one or more zoogeographic regions between central California and Washington. Density changes were far more variable in time and space, but in all regions, density declines were most often associated with large-bodied rockfish. We next estimated the impact of size and density changes on energy consumption and fecundity in a five-species rockfish assemblage that includes bocaccio (S. paucispinis), a large-bodied, overfished species. Indexes of both consumption and fecundity by the assemblage increased in the southern portion of the study area between 1980 and 2001 but decreased in the northern portion. Allocation of energy and reproductive potential within the assemblage shifted dramatically: relative to bocaccio, total energy consumption and fecundity indexes for the other four species increased by orders of magnitude from 1980 to 2001. These changes in community structure may affect the ability of bocaccio and other large rockfish species to recover from overfishing, especially in light of long-term declines in zooplankton production that may also be affecting rockfish size and production. Addressing these issues may require a regional, multispecies management approach.     

Habitat selection increases the isolating effect of habitat fragmentation on the gopher tortoise

Habitat selection requires choice, which differentiates it from habitat use, and choice, in turn, is dependent upon the responses of organisms to the environmental, social, and other cues that they perceive. Habitat selection by the gopher tortoise (Gopherus polyphemus) was investigated by translocating tortoises and monitoring their movements within two sites in central Florida. The first site supported a stable preponderance of high-quality habitat, and tortoises avoided areas with a dense tree canopy cover caused by fire exclusion. The second site was badly invaded by an introduced weed, and tortoises avoided areas where the weed had formed a dense monoculture. At both sites, individuals appeared to be responding to visual cues to avoid areas that were relatively dark. In landscapes with relatively large amounts of high-quality habitat, this avoidance behavior serves the gopher tortoise well by keeping individuals within the dominant habitat type. In degraded areas, high-quality habitat often becomes increasingly uncommon, and the avoidance behavior exhibited by the tortoises will result in individuals becoming confined to small patches, causing a significant reduction in fitness and hence questioning their long-term survival in such areas. The results from our study show that in order to maintain viable tortoise populations in areas increasingly subjected to human fragmentation and degradation, it is crucial not only to suppress tree canopy cover continually and prevent invasion by exotic weeds, but also to be mindful that the avoidance behavior of the gopher tortoise could prevent individuals from fully occupying a high-quality habitat in response to restoration and management efforts.     

Impact of sewage on the distribution,abundance and community structure of rocky intertidal macro-organisms

The biological effects of a low-volume domestic sewage discharge were studied near Wilson Cove, San Clemente Island, California (USA), from February to June, 1972. There were fewer species and less cover near the outfall (7 macro-invertebrates, 17.6%; 13 macrophytes, 91.7%) than in nearby “unpolluted” control areas (9 macroinvertebrates, 9.2%; 30 macrophytes, 103.4%). The outfall biota was less diverse than that of the controls, as shown by 5 different diversity indices. A great reduction in community stratification (spatial heterogeneity) and, hence, community complexity occurred near the outfall; this reduction in stratification was primarilly due to the absence of Egregia laevigata, Halidrys dioica, Sargassum agardhianum and Phyllospadix torreyi. These were replaced in the mid-intertidal near the outfall by a low turf of blue-green algae, Ulva californica, Gelidium pusillum and small Pterocladia capillacea, and in the lower intertidal by Serpulorbis squamigerus covered with Corallina officinalis Var. chilensis. A statistically-based determination of assemblages or groups of organisms (i.e., cluster analysis) revealed 3 discrete outfall and 3 discrete control area groups; 3 assemblages contained samples from both areas. The distributional patterns of these groups indicate that near the outfall the degree of dilution of discharged sewage is more important in regulating zonation than is tidal height. The enhancement of the suspension feeder Serpulorbis squamigerus and the omnivores Ligia occidentalis, Pachygrapsus crassipes and Anthopleura elegantissima in the outer fringe of the outfall plume hypothetically is due to their ability to utilize sewage as a food source. A critical effect of the outfall may be to decrease environmental stability thereby favoring rapid-colonizers and more sewage-tolerant organisms. The outfall macrophytes were characterized by relatively higher net primary productivities, smaller growth forms, simpler and shorter life histories, and most were components of early successional stages.     

Extra-pair paternity and the opportunity for sexual selection in a socially monogamous bird (Dendroica petechia)

We used multi-locus DNA fingerprinting to characterise the genetic mating system of the socially monogamous yellow warbler (Dendroica petechia). Over 2 years there were no instances of brood parasitism, but 59% of families (n = 90) contained extrapair sired young and 37% of offspring (n = 355) were of extra-pair paternity. Most hypotheses for extra-pair mating in monogamous species assume a paternity benefit to extra-pair sires, and focus on the benefit(s) to females. However, the assumption of male benefit has been little tested. Among yellow warblers, known extra-pair sires were just as likely to be cuckolded as any male in the population, and there was at least one reciprocal exchange of extra-pair paternity. Nevertheless, among known extra-pair sires, the paternity gains from extra-pair paternity were, on average, greater than the losses in their own families. These results show there is a paternity benefit to certain males. However, the benefit is not absolute but relative and therefore more difficult to measure. The results also suggest that patterns of extra-pair fertilisation are not determined by female choice alone. Most confirmed extra-pair mates were territorial neighbours, but some resided as far as three territories apart, and greater spatial separation was implied in other cases. Thus, the opportunity for extra-pair mating is great. We estimate that as a result of extra-pair fertilisations, variance in male mating success is increased somewhere between 3-fold and 15-fold over that which would result from within-pair reproduction alone. These findings affirm the potential importance of extra-pair reproduction for sexual selection in monogamous species and they support earlier suggestions that extra-territorial forays by male yellow warblers are for the purpose of extra-pair mating.     

Patterns of abundance, population size structure and microhabitat usage of Paracentrotus lividus (Echinodermata: Echinoidea) in SW Portugal and NW Italy

In coastal habitats, wave exposure influences several aspects of the life history of marine organisms. Here, we assess how hydrodynamic conditions can generate variation in density, size structure and microhabitat usage of Paracentrotus lividus and whether these effects are consistent between regions that are markedly different for oceanic climate, such as the coasts of SW Portugal and NW Italy. The abundance of P. lividus was ~4 times higher in SW Portugal than in NW Italy, but within each region, there was no effect of wave exposure. In SW Portugal, higher urchin abundances were found at shallower depths, while no effect of depth on urchin abundance emerged in NW Italy. Most of the variation in urchin abundance occurred at small spatial scales (metres), and our results suggest that habitat complexity, that is, the presence of cracks and crevices, is an important determinant of patterns of distribution of this species. The population in NW Italy was characterized by a unimodal size distribution, with a higher proportion of medium-sized individuals. In contrast, in SW Portugal, smaller individuals represented a large proportion of the populations. Size structure varied between exposed and sheltered habitats in SW Portugal, suggesting that the proportion of individuals from different size cohorts may vary along wave-exposure gradients as a result of direct or indirect effects of hydrodynamic forces. In SW Portugal, most urchins occurred in burrows, while in NW Italy, urchins were mainly observed in crevices. These results suggest that creating/occupying burrows might be an adaptive behaviour that allows sea urchins to better withstand stressful hydrodynamic conditions and, therefore, are more common on exposed Atlantic coasts. Overall, our study suggests that the effects of hydrodynamic forces on sea urchin populations are context dependent and vary according to background oceanic climate.     

Vertical distribution,abundance and community structure of oncaeid copepods in the Oyashio region,western subarctic Pacific

The vertical distribution, abundance and community structure of oncaeid copepods were investigated in the Oyashio region, western subarctic Pacific. Samples were collected with a 0.10 mm mesh closing type net from five discrete layers down to a maximum depth of 2,000 m in September and December 1996 and in April and October 1997. The copepods were widely distributed from epipelagic to bathypelagic zones, and showed prominent peaks of abundance above the thermocline and/or between 250 and 1,000 m depth. Standing stocks of total oncaeid copepods ranged from 1.5 to 2.5×10⁵ inds m^–2 at 0–2,000 m in the water column, which are the same order of magnitudes as those reported in tropical, subtropical and polar regions by previous workers. A total of 38 species and two forms belonging to the genera Oncaea, Triconia, Spinoncaea, Conaea and Epicalymma, and two provisionally classified species of the family Oncaeidae were identified in this study. Of these, 14 species have already been recorded from the eastern subarctic Pacific. Several warm-water species were also found in December 1996 and/or October 1997, when the effect of warm-core rings originating from the Kuroshio Current was evident in the epipelagic zone. Dominant species throughout the whole water column were T. borealis, T. canadensis, O. grossa, O. parila, O. rimula, O. lacinia, Epicalymma spp. and Oncaea sp. A, and these eight species together always made up about half of the total oncaeid numbers. Community analysis revealed that species composition in epipelagic and upper mesopelagic zones varied temporally due to the changes of hydrographic conditions, whereas those in lower mesopelagic and bathypelagic zones were relatively stable, reflecting the constancy of deep environments.Communicated by O. Kinne, Oldendorf/Luhe     

Habitat selection of two congeneric bivalves,Cardium edule and C. glaucum in sympatric and allopatric populations

Habitat selection of two bivalve species, Cardium edule (L.) and C. glaucum (Bruguière) is described for sympatric and allopatric populations of both sibling species. It is demonstrated that C. edule grows faster than C. glaucum and is absent from localities with periodic low food-availability. The ability to burrow of C. glaucum is low compared with that of C. edule; the former species does not occur in very loose, well-sorted sand near the coast. Earlier studies which suggest that tidal range determine the species distribution could not be confirmed.     

Distribution patterns,abundance and population dynamics of the euphausiidsNyctiphanes capensis andEuphausia hanseni in the northern Benguela upwelling system

Distribution patterns, population structure and biomass of the euphausiidsNyctiphanes capensis andEuphausia hanseni were examined off the coast of Namibia, southwest Africa, in relation to temperature, depth and season, from data collected on nine surveys from September 1982 to March 1984. High densities ofN. capensis were found in the shallow coastal waters (<200 m), with the biomass of adults ranging from 675 to 5 706 mg dry wt m^–2. For adultE. hanseni, the biomass was an order of magnitude lower, ranging from 65 to 505 mg dry wt m^–2, with most specimens occurring over the shelf break at depths of 200 to 1000 m. These distribution patterns remained relatively constant throughout the year, despite seasonal differences in upwelling events. Both species displayed continuous breeding, with 43 to 82% of the adult femaleE. hanseni being fertilized, while a much lower proportion ofN. capensis females were reproductively active (0.5 to 26%). Different breeding strategies were adopted by these two euphausiid species, withE. hanseni producing frequent broods (14.8 broods in 6 mo) consisting of relatively large eggs which are released into the sea, andN. capensis exhibiting a lower frequency of spawning, with broods consisting of large numbers of relatively small eggs, protected by a brood pouch. These strategies enable both species to maintain high densities throughout the year in a fluctuating physical environment. Growth rate estimated from size-frequency distributions were 0.003 to 0.063 mm d^–1 forN. capensis and 0.077 to 0.083 mm d^–1 forE. hanseni, suggesting an adult lifespan of approximately 6 mo for both species. Maximum sizes were attained in September, withN. capensis reaching a total length of 21 mm (in contrast to all previous studies onN. capensis, where the maximum size recorded was only 13 mm total length) andE. hanseni a total length of 33 mm.