A new hypothesis concerning the nature of small pelagic fish clusters: An individual-based modelling study of Sardinella aurita dynamics off West Africa

Coastal populations of small pelagic fish display nested aggregation levels. Above the level of the school structure, clusters are observed the nature of which has not been definitively determined. We hypothesized that these clusters corresponded to a materialisation of the microcohorts originating from successive spawnings of fish populations in their vital domain.A candidate individual-based model was developed to investigate this hypothesis. This model is based on pattern-oriented modelling of a concrete documented case: the dynamics of the round sardinella (Sardinella aurita) population living off the West African coasts and subject to environmental fluctuations caused by seasonal upwelling. The simulated agents were round sardinella microcohorts situated and moving in a discretised physical environment. The combined effects of environmental forcing (temperature, wind, retention) and inner biological dynamics (reproduction, growth and mortality, competition) condition the dynamics of this population.The modelled behaviour generated realistic dynamic patterns (population distribution, spawning zones, periods and plasticity, biomass fluctuations), which were obtained simultaneously and successfully compared with observations. The steady-state number of microcohorts obtained after simulation convergence was similar to the number of clusters observed in situ in this area for this population.The realism and diversity of the patterns simultaneously simulated suggested the cluster-microcohort equivalence hypothesis as a candidate framework accounting for the origin of the clusters observed in situ. Within this preliminary exploration, we discuss the consistency of the hypothesis and the accuracy of the model. If the correspondence between clusters and microcohorts proves to be real, it may be transient and progressively modified by other environmental factors. If stable over time, as simulated in the model, the number of observed clusters should be related to the number of spawning events in the species’ lifetime.     

Mitochondrial DNA variation and population structure in the spiny lobster Panulirus argus

Adult spiny lobsters (Panulirus argus) were collected from nine locations throughout the tropical and subtropical northwest Atlantic Ocean and examined for mitochondrial DNA (mtDNA) variation. 187 different mtDNA haplotypes were observed among the 259 lobsters sampled. Haplotype diversity was calulated to be 0.986 and mean nucleotide sequence-diversity was estimated to be 1.44%; both of these values are among the highest reported values for a marine species. Analysis of molecular variance (AMOVA) and phenetic clustering both failed to reveal any evidence of genetic structure within and among populations of P. argus. The present data are consistent with high levels of gene flow among populations of P. argus resulting from an extended planktonic larval stage and strong prevailing ocean currents.     

Results of the UNDP (SF)/FAO regional fisheries survey in West Africa. Report No. 1. Deep-scattering Sardinella aurita off Mauritania

Scattering layers beyond the Continental Shelf, in deep water, were often observed during cruises with the charter vessel of the Regional Fisheries Survey in West Africa. The layers were located at depths of 100 to 350 m during daytime; at night, they generally ascended to the surface. They consisted mainly of myctophids and small squids. In this report, results of first observations on deep-scattering layers of Sardinella aurita (Cuv. et Val.) are given. The layers occurred off Mauritania during the months of hydrological transition, characterized by low water-temperatures. In these months, S. aurita were rarely caught over the shelf. Apparently, they migrate north and south in deep water, and were not detected by commercial vessels. The environmental conditions and the difficulties for the industrial fishery during this transitional period are briefly described. the temporal and spatial occurrence of deep-scattering S. aurita is given, and diurnal vertical migrations are shown, together with examples of echo records. Results of fishing operations made in the layer, and some biological observations on S. aurita are presented.     

Behavioral responses of territorial red squirrels to natural and experimental variation in population density

The relative scarcity of studies at the intersection of behavioral and population ecology is surprising given the presumed importance of behavior in density-dependent population regulation. Here we tested whether North American red squirrels (Tamiasciurus hudsonicus) adjust their behavior in response to local population density and whether they use rates of territorial vocalizations in their local neighborhood to assess density. We examined these relationships using 18?years of live trapping and 20?years of behavioral data that were collected across natural variation in local population density. To disentangle the effects of population density on behavior from those due to changes in per capita food abundance or changes in the frequency of antagonistic interactions with neighbors, we also experimentally manipulated population density with long-term food supplementation as well as perceived population density with long-term playbacks of territorial vocalizations. The frequency with which squirrels emitted territorial vocalizations was positively associated with local population density. In contrast, antagonistic physical interactions observed between squirrels and territorial intrusions were rare and the frequency of intrusions was weakly and negatively, not positively, associated with population density. Squirrels experiencing naturally and experimentally high density conditions spent less time in the nest and feeding but more time being vigilant. Similar density-dependent changes in behavior were observed in response to our manipulations of perceived population density, indicating that vocalization rates and not physical interactions or food abundance were the mechanism by which squirrels assessed and responded behaviorally to changes in local density.     

Population structure,northern range limit,and recruitment variation in the intertidal limpet Lottia scabra

The northern range limit of the intertidal limpet Lottia scabra is Cape Arago, Oregon (43°N), where adult survival is excellent, the population is small (<300), and recruitment is low; the range limit may be set by limited recruitment. Between June 2012 and March 2013, 25 sites from the middle of the species range (33°N) to Cape Arago were sampled and population size frequency distributions, densities, and nearest neighbor distances were compared to the amount of rocky and sandy shore and kelp bed size. North and south of 37°N, the densities of new recruits averaged 22 and 86 m^?2, respectively. This shift was associated with the range limit of Macrocystis pyrifera kelp beds; we hypothesize that slower currents in M. pyrifera beds may limit larval dispersal leading to higher recruitment. North and south of 40°N, adult density averaged <1 and 458 m^?2, respectively, with the species absent from many sites to the north. This shift was associated with a sharp drop in the amount of rocky shoreline and an increase in uninhabitable sandy shore. Near the northern range limit, >80 % of the individuals were solitary and may be unable to spawn successfully. Recruitment at Cape Arago was infrequent and likely due to self-recruitment. This study suggests that the range limit was set by the absence of M. pyrifera and too little rocky shore leading to high larval wastage, low settlement, low population densities, and, due to an Allee effect, very small effective population sizes.     

Estimating digestion time in gelatinous predators: a methodological comparison with the scyphomedusa Aurelia aurita

In order to quantify the trophic impact of gelatinous predators, digestion time estimates are commonly applied to counts of prey in the guts. Three primary approaches are used, the Manual-feeding, Natural-feeding and Steady-state methods; these differ in methodology and their underlying assumptions. The criteria used to define the end-point of digestion, and the resolution at which digestion progress is observed, also vary across studies. To understand the impact of such differences, we estimate digestion times of the scyphomedusa Aurelia aurita fed adult females of the copepod Acartia tonsa using these various approaches. We find ~fourfold differences which can be attributed to bias towards the slowest rates of digestion by some end-point criteria, and overestimation from low observation resolution. Artificial manipulation and the degree to which swimming and feeding behaviour are natural may also influence estimates. We provide recommendations for those quantifying digestion times of Aurelia aurita medusae and gelatinous predators.     

Population genetic variation in the New Zealand greenshell mussel: locus-dependent conflicting signals of weak structure and high gene flow balanced against pronounced structure and high self-recruitment

A pronounced north–south multitaxon genetic discontinuity occurs in central New Zealand (NZ). Polymorphic microsatellite markers have been used to test the location and structure of this discontinuity in the endemic greenshell mussel, Perna canaliculus. Nine neutral loci revealed limited evidence of genetic structure, but one outlier locus (Pcan1-27) which may be under selection provided evidence of the discontinuity in central NZ. Whilst the limited multilocus evidence of structure is assumed to result from high levels of gene flow among populations of this continuously distributed species, assignment tests indicated high to very high mean levels of self-recruitment within the 14 populations and the north and south regions. The nine neutral loci were unable to provide further clarification as to the geographic location of the discontinuity, whereas the Pcan1-27 locus was particularly informative. These results highlight a tension between limited evidence of genetic structure and presumptive high gene flow among populations versus high levels of self-recruitment and pronounced structure depending on microsatellite loci and analyses in question. Evidence from all 10 loci indicates that the genetic discontinuity is maintained by high levels of self-recruitment, and evidence from Pcan1-27 suggests that selection may also be important in explaining the existence of the discontinuity.     

Population genetics of the blue crabCallinectes sapidus: modest population structuring in a background of high gene flow

To determine the genetic population structure of blue crabs (Callinectes sapidus Rathbun), electrophoretic allozyme analysis was performed on 750 individuals collected from 16 nearshore locations ranging from New York to Texas, USA. Twenty enzymes and non-enzymatic proteins coded by 31 presumptive loci were examined. Twenty-two loci were either monomorphic or polymorphic at less than theP ₉₅ level; alleles for these polymorphic loci were geographically dispersed. Allele frequencies for three of the remaining polymorphic loci were homogeneous over all populations, as were levels of polymorphism and heterozygosity. Phenograms generated by the UPGMA (unweighted pair-group method using arithmetic averages) and distance Wagner methods exhibited no geographic pattern in the clustering of populations. Estimates ofN _em (effective number of migrants per generation between populations) indicated substantial gene flow, with aalues sufficiently high to infer panmixia between all blue crab populations from New York to Texas. However, despite this high level of gene flow, two striking patterns of geographic differentiation occurred: genetic patchiness and clinal variation. Allele frequencies atEST-2, GP-1, IDHP-2, DPEP-1, DPEP-2, andTPEP exhibited genetic patchiness on local and range-wide geographic scales, and allele frequencies atEST-2 varied temporally. Genetic patchiness in blue crabs is likely to be the result of the pre-settlement formation and subsequent settlement of genetically heterogeneous patches of larvae; allele frequencies of those larval patches may then be further modified through ontogeny by localized selection. In the Atlantic Ocean, a regional latitudinal cline ofEST-2 allele frequencies was superimposed on the range-wide genetic patchiness exhibited by that locus. This pattern against a background of high gene flow is highly likely to be maintained by selection. In estuaries along the Atlantic Ocean coast, a combination of low adult long-distance migration and a high retention rate of locally spawned larvae could serve to segregate populations and allow for the development of the geographic cline inEST-2. The Gulf of Mexico showed no apparent cline, perhaps due to long-distance migration of females in some regions of the Gulf, or to masking by genetic patchiness. These results emphasize the importance of both ecological and evolutionary time scales and structuring mechanisms in determining genetic population structure.     

Short-term temporal variation in population structure of two harpacticoid copepods,Zausodes arenicolus and Paradactylopodia brevicornis

Aspects of the population biology of two harpacticoid copepod species [Zausodes arenicolus Wilson and Paradactylopodia brevicornis (Claus)] living in Tampa Bay, Florida, USA, were studied on time scales of hours to days during November 1983. Fluctuations in population agestructure, adult sex-ratio, and proportion of ovigerous females were determined for these two species in a control plot and in an experimental plot defaunated by raking. For both species, immigrants into the defaunated plot differed significantly from conspecifics in the control plot in all characteristics examined. The disturbed plot contained a larger proportion of young juveniles of both species compared to the unmanipulated site after 4 h but not after 8 h. In Z. arenicolus, adult males returned to the experimental plot more quickly than adult females. For both species, rapid and significant changes in population age-structure, adult sex-ratio, and the proportion of adult females with eggs occurred within the control plot. It is suggested that this variation was too large and occurred too rapidly to be accounted for solely by in situ processes in the absence of large-scale immigration and/or emigration of individuals. Potential implications of these results for population studies of harpacticoid copepods and other benthic invertebrates are discussed.     

Morphological variation in a population of grey-crowned babblers: Correlations with variables affecting social behavior

Summary Variation in weight, wing length and bill length in a population of grey-crowned babblers is influenced primarily by sex and age, but correlations with size of social unit, with reproductive success, and with vegetation are also detectable (Table 1). The latter correlations vary with sex, age, and status as helper or breeder. Differential wear according to behavioral role, competition for status, incubation, and inheritance are discussed as possible causal mechanisms. Helpers were not detectably smaller in any dimension than breeders of the same age and sex. Male and female non-breeding helpers differ in patterns of morphological correlation, suggesting that they have different behavioral roles. Breeding males have a unique pattern of morphological correlation, suggesting that their foraging behavior differs from breeding females and non-breeders.     

Mitochondrial DNA variation in Eritrean hamadryas baboons (Papio hamadryas hamadryas): life history influences population genetic structure

The hamadryas baboon, Papio hamadryas hamadryas, represents a rare exception from the pattern of female philopatry and male-biased dispersal predominant in mammals including primates. To elucidate the possible consequences of the dispersal pattern on the population genetic structure of hamadryas baboons, we sequenced the maternally transmitted mitochondrial hypervariable region I of 74 individuals from ten sampling locations in different ecogeographic zones of Eritrea. To this end, individual fecal samples were collected at sleeping cliffs. Upon comparing the individual sequences by means of phylogenetic tree reconstructions and AMOVA, we could not detect a population genetic structure corresponding to a geographic pattern. Tree reconstructions revealed the existence of two profoundly different lineages both present at most of the sampling locations. These findings and Mantel correlations of genetic distances and the frequency of shared haplotypes to geographic distances point to the presence of female dispersal. Female-mediated gene flow is detectable over geographic distances exceeding those between neighboring subpopulations. Our study therefore corroborates local behavioral observations on a broad geographic scale. After inclusion of geographically closely situated olive baboons, P. h. anubis, in the analyses, all anubis sequences fell within one hamadryas clade. Possible scenarios leading to this situation including long-term hybridization processes are discussed.     

Microsatellite variation, effective population size, and population genetic structure of vermilion snapper, Rhomboplites aurorubens, off the southeastern USA

Vermilion snapper (Rhomboplites aurorubens) were collected from four sites off the Atlantic coast of the USA and one site in the Gulf of Mexico to evaluate effective population size and genetic stock structure. Previous studies had suggested geographic variation in the ratio of males to females, so this population characteristic was explored in conjunction with the genetic analysis. Sex ratio varied greatly among the five sample sites; males comprised 57% of samples in the Gulf of Mexico, while within the South Atlantic Bight they comprised between 36% (Morehead City, North Carolina) and 53% (Carolina Beach, North Carolina) of samples. No clear geographic trends in the sex ratio emerged; instead, it was found to vary with fish length, the percentage of males decreasing with increasing size. Allelic variation assessed at seven dinucleotide microsatellite loci was large; gene diversities ranged from 0.43 to 0.95 and allelic counts from 7 to 39. Estimates of the effective population size ranged from 24 500 (based on the infinite-alleles model) to 150 500 (based on the stepwise-mutation model). There was evidence for excess homozygosity within samples: estimates of F _IS (the correlation of alleles within individuals) ranged from 0.01 to 0.03 among the seven loci, and three estimates were significantly greater than zero. Differentiation among localities was very weak, as estimates of F _ST (the correlation of alleles within populations) were on the order of 0.001 to 0.002 and genetic distance estimates between localities were not related to geographic distances. This suggested that vermilion snapper in the South Atlantic Bight (Cape Hatteras, North Carolina to Cape Canaveral, Florida) and Gulf of Mexico are likely to consist of one genetic stock. Despite the overall homogeneity, there were indications of a temporally dynamic local structure that would bear further examination. Received: 6 July 1998 / Accepted: 9 February 1999     

Sources of variance in a female fertility signal: exaggerated estrous swellings in a natural population of baboons

Signals of fertility in female animals are of increasing interest to evolutionary biologists, a development that coincides with increasing interest in male mate choice and the potential for female traits to evolve under sexual selection. We characterized variation in size of an exaggerated female fertility signal in baboons and investigated the sources of that variance. The number of sexual cycles that a female had experienced after her most recent pregnancy (“cycles since resumption”) was the strongest predictor of swelling size. Furthermore, the relationship between cycles since resumption and swelling size was most evident during rainy periods and was not evident during times of drought. Finally, we found significant differences in swelling size between individual females; these differences endured across cycles (i.e., were not explained by variation within individuals) and persisted in spite of ecological effects. This study is the first to provide conclusive evidence of significant variation in swelling size between female primates (controlling for cycles since resumption) and to demonstrate that ecological constraints influence variation in this signal of fertility.     

Infaunal benthos of a natural petroleum seep: Study of community structure

At 8 wk intervals, from December 1975 through March 1978, we took a census of the infaunal benthic invertebrates at a natural petroleum seep near Santa Barbara, California, USA and at an area nearby where no fresh petroleum occurred in the sediments. Both sites had in common 72% of the populations representing over 90% of the individuals, strongly suggesting that the two sites are part of the same community. At the petroleum seep there were higher densities of individuals in many populations (60%), but no dramatic difference in diversity (Shannon-Wiener or dominance-diversity). The seep populations tended to be more variable from one sampling period to the next, possibly due, at least in part, to large larval settlements there. The most abundant populations at the seep site were dominated by deposit feeders (14 of the 15 species considered), especially oligochaetes which are extremely rare at the comparison site. The discussion includes a hypothesis of trophic enrichment of the seep populations by bacterial growth stimulated by fresh petroleum.