Density dependence in marine fish populations revealed at small and large spatial scales

Experimental manipulation of population density has frequently been used to demonstrate demographic density dependence. However, such studies are usually small scale and typically provide evidence of spatial (within-generation) density dependence. It is often unclear whether small-scale, experimental tests of spatial density dependence will accurately describe temporal (between-generation) density dependence required for population regulation. Understanding the mechanisms generating density dependence may provide a link between spatial experiments and temporal regulation of populations. In this study, I manipulated the density of recently settled kelp rockfish (Sebastes atrovirens) in both the presence and absence of predators to test for density-dependent mortality and whether predation was the mechanism responsible. I also examined mortality of rockfish cohorts within kelp beds throughout central California to evaluate temporal (between-generation) density dependence in mortality. Experiments suggested that short-term behavioral responses of predators and/or a shortage of prey refuges caused spatial density dependence. Temporal density dependence in mortality was also detected at larger spatial scales for several species of rockfish. It is likely that short-term responses of predators generated both spatial and temporal density dependence in mortality. Spatial experiments that describe the causal mechanisms generating density dependence may therefore be valuable in describing temporal density dependence and population regulation.     

Escape speeds of marine fish larvae during early development and starvation

Response rates to tactile stimulation and subsequent escape speeds were measured using a video-recording system during early development and starvation of fish larvae. The species studied included the yolk-sac larvae of Clyde and Baltic herring (Clupea harengus L.), cod (Gadus morhua L.), flounder (Platichthys flesus L.) and older larvae of Clyde herring. The proportion of larvae responding (response rate) was initially about 20 to 25% in herring and 35 to 40% in cod and flounder using a probe, but about 70 to 80% using the sucking action of a pipette in all species except flounder. Both response rates and escape speeds (mean and maximum) tended to peak 1 to 2 d before the PNR (point-of-no-return, when 50% of larvae are too weak to feed), then decreased slowly during further starvation. An inter-species comparison showed that the highest recorded mean escape speeds (measured over a period of 200 ms) and highest maximum escape speeds (over 20 ms) ranged from 5.7 to 8.6 BL/s (body lengths/s) and 12.1 to 16.1 BL/s, respectively. The larvae made directional responses away from the stimulus only when they developed and reached the feeding stage.     

Osmoregulation, nutritional effects and buoyancy of marine larval fish: a bioassay for assessing density changes during the earliest life-history stages

It has been hypothesized that marine fish larvae in the advanced stages of starvation would show increased density (ρ = mass volume⁻¹) from water loss due to osmoregulation failure. Changes in larval buoyancy are currently attributed to swim bladder regulation and protein synthesis or catabolism. Osmoregulation-related changes in density is an alternative mechanism, the importance of which remains untested in the laboratory and the influence of which on vertical distributions is unknown. We provide evidence that loss of osmotic control is a plausible mechanism for increased density of larval cod (Gadus morhua L.). Furthermore, our results show that this mechanism is not restricted to larvae in the advanced stages of starvation. “Relative” larval densities are estimated using a modified density gradient. We use a gravimetric method to separate the effects of nutrition from osmoregulation failure. We assessed the importance of sampling strata on estimates of larval density. Proportional sampling within three depth strata (stratified sample) produced the least biased method for determining the “average” density of a population of larvae in laboratory culture. Larvae sampled from the bottom third of the culture tank were significantly more dense then those sampled from the surface. This was true for larvae of all ages. The average change in density from hatching till death from starvation for larvae sampled in the surface stratum was nominal (Δρ = 5.0 × 10⁻⁴ g cm⁻³), while the change for those sampled from the bottom stratum was large (Δρ = 3.8 × 10⁻³ g cm⁻³). These large density differences suggest that larvae sampled from the bottom stratum were either osmotically stressed or were facultatively changing their density via regulatory pathways. Preliminary observations suggest that vitality is lower amongst those larvae which are sampled near the bottom. The small change in average density of larvae sampled from the surface stratum was due to starvation. The density differences we observed between “osmotically stressed” and “starving” larvae could readily have been misconstrued as differences in feeding and growth experienced by individual larvae. The potential bias of increased density from osmoregulation failure must be considered as a factor in experimental designs developed to assess the effect of fed and starved treatments on buoyancy for larvae of all ages. The simple bioassay we describe may prove useful both as a means of assessing larval condition and as a mechanism for evaluating factors affecting larval vertical distributions in the field. Received: 13 January 1997 / Accepted: 3 February 1997     

Density dependence vs. independence, and irregular population dynamics of a swallow-wort fruit fly

Although most long-term studies of consumer-resource (e.g., predator-prey) interactions select species showing cyclic population dynamics, strong consumer-resource interactions can also produce irregular, noncyclic dynamics. Here, we present a case in which a seed predator, the tephritid fruit fly Euphranta connexa, shows fluctuations in density of more than two orders of magnitude over a 22-year period. To explain these fluctuations, we analyzed a stage-specific data set to quantify the density-dependent and density-independent components of larval survivorship and realized fecundity. Both larval survivorship and realized fecundity were strongly density dependent. Larval survivorship dropped from 0.62 at low larval density to 0.081 at high larval density, whereas fecundity dropped from 84.3 to 0.32 eggs per individual, more than a 100-fold decrease. We divided density-independent variation in E. connexa population dynamics into components for variability in (1) larval survivorship, (2) realized fecundity, and (3) annual fruit abundance. Of these components, 96% of the density-independent variance in per capita population growth rates was caused by fluctuations in fruit abundance. This highlights the importance of the strong consumer-resource interactions in driving fluctuations in E. connexa abundance. It also demonstrates that E. connexa dynamics are remarkably simple, and aside from the 4% of unexplained variance in per capita population growth rates, our understanding of E. connexa dynamics is remarkably complete.     

Estimating the functional form for the density dependence from life history data

Two contrasting approaches to the analysis of population dynamics are currently popular: demographic approaches where the associations between demographic rates and statistics summarizing the population dynamics are identified; and time series approaches where the associations between population dynamics, population density, and environmental covariates are investigated. In this paper, we develop an approach to combine these methods and apply it to detailed data from Soay sheep (Ovis aries). We examine how density dependence and climate contribute to fluctuations in population size via age- and sex-specific demographic rates, and how fluctuations in demographic structure influence population dynamics. Density dependence contributes most, followed by climatic variation, age structure fluctuations and interactions between density and climate. We then simplify the density-dependent, stochastic, age-structured demographic model and derive a new phenomenological time series which captures the dynamics better than previously selected functions. The simple method we develop has potential to provide substantial insight into the relative contributions of population and individual-level processes to the dynamics of populations in stochastic environments.     

Parasitism and a shortage of refuges jointly mediate the strength of density dependence in a reef fish

Various predator-prey, host-pathogen, and competitive interactions can combine to cause density dependence in population growth. Despite this possibility, most empirical tests for density-dependent interactions have focused on single mechanisms. Here we tested the hypothesis that two mechanisms of density dependence, parasitism and a shortage of refuges, jointly influence the strength of density-dependent mortality. We used mark-recapture analysis to estimate mortality of the host species, the bridled goby (Coryphopterus glaucofraenum). Sixty-three marked gobies were infected with a copepod gill parasite (Pharodes tortugensis), and 188 were uninfected. We used the spatial scale at which gobies were clustered naturally (approximately 4 m2) as an ecologically relevant neighborhood and measured goby density and the availability of refuges from predators within each goby's neighborhood. Goby survival generally declined with increasing density, and this decline was steeper for gobies with access to few refuges than for gobies in neighborhoods where refuges were common. The negative effects of high density and refuge shortage were also more severe for parasitized gobies than for gobies free of parasites. This parasite has characteristics typical of emerging diseases and appears to have altered the strength of a preexisting density-dependent interaction.     

Density and distribution patterns of the temperate marine fish Cheilodactylus spectabilis (Cheilodactylidae) in a reef environment

Cheilodactylus spectabilis (Hutton) is common over shallow reefs in north-eastern New Zealand. Replicated transect between-area differences in density and in size frequency. C. spectabilis is not nocturnally active. The between-area differences in density and size frequency remained constant over daylight hours, demonstrating that C. spectabilis does not undergo any systematic feeding migrations. Topographic complexity was shown to have a significant positive influence on fish density. Mean size was related to water depth at 5 localities investigated, with small (<200 mm standard length, SL) individuals being restricted to shallow water. All sizes of C. spectabilis examined had fed on small invertebrates, with gammarid amphipods predominating. No evidence of size-related differentiation in feeding patterns was observed, although small individuals spent significantly more time feeding than large ones. Small (<200 mm SL) individuals occupied feeding areas and shelter sites from which they excluded other small C. spectabilis. Large individuals showed no site-associated aggression, covered a greater area during daily movements and overlapped with other large and small C. spectabilis while feeding. No evidence of dawn or dusk peaks of feeding activity or movement was collected.     

Modelling effects of time-variable exposure to the pyrethroid beta-cyfluthrin on rainbow trout early life stages

Background

Available literature and regulatory studies show that the severity of effects of beta-cyfluthrin (a synthetic pyrethroid) on fish is influenced by the magnitude and duration of exposure. To investigate how the exposure pattern to beta-cyfluthrin (constant vs peak) may influence the response of the fish, we used a mechanistic effect model to predict the survival and growth of the rainbow trout over its early life stages (i.e. egg, alevin and swim-up fry). We parameterized a toxicokinetic–toxicodynamic (TKTD) module in combination with a dynamic energy budget model enabling us to describe uptake and elimination, as well as to predict the threshold concentration for survival and sublethal effects (feeding behaviour and growth). This effect model was calibrated using data from an early life stage experiment where trout was exposed to a constant concentration of cyfluthrin. The model was validated by comparing model predictions to independent data from a pulsed-exposure study with early life stages of rainbow trout.

Results

The co-occurrence of effects on behaviour and growth raised the possibility that these were interrelated, i.e. impairment of feeding behaviour may have led to reduced food intake and slower growth. We, therefore, included ‘effect on feeding’ as mode of action in the TKTD module. At higher concentrations, the constant exposure led to death. The model was able to adequately capture this effect pattern in the calibration. The model was able to adequately predict the response of fish eggs, alevins and swim-up fry, from both the qualitative (response pattern) and quantitative points of view.

Conclusions

Since the model was successfully validated, it can be used to predict survival and growth of early life stages under various realistic time-variable exposure profiles (e.g. profiles from FOCUS surface water modelling) of beta-cyfluthrin.

     

Effect of thermal stress and total residual chlorination on early life stages of the mummichog Fundulus heteroclitus

Effects of simultaneous short-term (7.5 to 60 min) thermal stress (24° to 34°C) and total residual chlorination (0.05 to 1.0 mg l^-1) on specific development stages of the mummichog Fundulus heteroclitus (Pisces: Cyprinodontidae), were investigated. For the embryonic stages, the total number of successfully hatched larvae was used as the criterion to measure effect. For the larval stages, survival 24 h after exposure was used. In the embryonic stages, temperature was the most important main variable. Only one embryonic stage (gastrula) was confounded by second-order interactions (temperature x duration of exposure x total residual chlorination). Both 0-day and 7-day-old larval stages showed significant higher-order interactions for all combinations of test parameters, suggesting the presence of synergistic effects of the three main experimental variables.Contribution No. 308 to the Gulf Breeze Environmental Research Laboratory. Contribution No. 183 to the Belle W. Baruch Institute for Marine Biology and Coastal Research.     

Respiration of Sepia officinalis during embryonic and early juvenile life

Adult Sepia officinalis L. were caught in June 1984, in the coastal waters of Wimereux (France). Deposition of the eggs took place in the seawater aquaria of the Station Marine. The oxygen consumption of S. officinalis was measured during embryonic and juvenile development. Aerobic metabolism occurs as soon as the early embryonic Stage 21. Oxygen diffuses through the initially thick egg shell; the oxygen level in the perivitelline liquid reaches a maximal value just before hatching (116.7±6.9 mm Hg). Hatchings display only a slight increase in oxygen consumption compared to embryos in the last stage of development. Respiration experiments with 40 d old juveniles showed that oxygen consumption increases with temperature, but is not affected by photoperiod. Experiments under increasing hypoxia revealed that S. officinalis juveniles are good regulators and maintain a constant oxygen consumption in the range of 4 to 7 mg O₂l^-1. Juveniles successfully recover from an hypoxic stress of 2 mg O₂l^-1 maintained for 1 h. This suggests that the respiratory pigments (pre-hemocyanins) of 40 d-old juveniles have a high oxygen affinity and/or that these juveniles have the ability to adapt to anaerobic conditions.     

Development and characterization of complex particles for delivery of amino acids to early marine fish larvae

Complex particles (CP), zein microbound particles and lipid-spray beads (LSB) were prepared and their performances were compared for delivering glycine and a mixture of free amino acids (FAA; alanine, glycine, leucine, serine and tyrosine) to early fish larvae. Measures of performances of microparticles included inclusion, encapsulation, retention and delivery efficiencies in addition to T50 (time to 50% retention) values. CP were prepared containing LSB and a defined dietary mixture that were bound together with zein. CP had significantly higher retention and delivery efficiencies for FAA compared to those of zein microbound particles. Free LSB had higher retention efficiencies for particulate glycine compared with CP, possibly due to differences in suspension characteristics. Free LSB clumped and floated when suspended in water, negatively affecting their acceptability by fish larvae; therefore, LSB should be incorporated into CP for more effective delivery of amino acids. There was a significant inverse correlation between retention efficiencies and solubilities of FAA encapsulated in CP. After 1 h of aqueous suspension, highest retention (44%) and delivery efficiencies (20.3 mg tyrosine g^–1 particle) were achieved with tyrosine. T50 values indicated 50% of the initial tyrosine in CP was still available after 36.7 min of suspension in water. Furthermore, CP can deliver FAA to marine fish larvae based on the results of feeding studies in which CP were digested by 3-day-old clownfish larvae; therefore, this particle type could be a valuable tool in studies of larval fish nutrition.Communicated by R.J. Thompson, St. Johns     

Effects of fish feeding by snorkellers on the density and size distribution of fishes in a Mediterranean marine protected area

Although there is a great deal of evidence to show that supplementary feeding by humans in terrestrial environments causes pronounced changes in the distribution and behaviour of wild animals, at present very little is known about the potential for such effects on marine fish. This study evaluated the consequences of feeding by snorkellers on fish assemblages in the no-take area of the Ustica Island marine protected area (MPA; western Mediterranean) by (1) determining if reef fish assemblage structure is affected in space and time by tourists feeding the fish; (2) assessing the effects of feeding on the abundance of the most common fish species; and (3) assessing the effects of feeding on the size structure of the two most numerically dominant ones. In particular, we hypothesised that both the abundance and the size structure of some fish species would increase at the study site following supplementary feeding, since the additional food provided by humans would make the site more appealing to them. Fish feeding influenced the fish assemblages within the Ustica MPA, and significant spatio-temporal changes occurred. While fish feeding appeared to have no effect on the ornate wrasse Thalassoma pavo, there was a noticeable increase in the number of Oblada melanura and Epinephelus marginatus in the impacted location after feeding. It is very likely that aggregations of fishes that evolve as a result of fish feeding by the public may have negative effects on local populations of fishes and invertebrates that make up their prey. Recreational use of coastal areas and MPAs is increasing elsewhere, making fish feeding a generalised human activity. Accurate information about its effect on the fish assemblage is essential to make responsible management decisions.Communicated by R. Cattaneo-Vietti, Genova     

Thermal tolerance of Strongylocentrotus purpuratus early life history stages: mortality, stress-induced gene expression and biogeographic patterns

In this study, we examined the differential thermal tolerance of Strongylocentrotus purpuratus early life history stages by comparing high temperature–induced mortality and the relative levels of the stress-induced gene, hsp70, between S. purpuratus embryos and larvae from adults collected throughout the species range. There was no significant difference between gastrulae and 4-arm plutei mortality from all sites examined. Furthermore, there was little variability in temperature tolerance across the biogeographic range as southern gastrulae and 4-arm plutei exhibited similar tolerances to northern individuals. Relative levels of hsp70 mRNA expression did not differ overall between the two developmental stages at each site. Across sites, all gastrulae and 4-arm plutei exhibited maximum hsp70 expression at approximately 25°C; however, the range of hsp70 expression was narrower in southern individuals, suggesting they are living closer to their upper thermal limit than northern individuals.     

Effects of the dinoflagellates Karlodinium veneficum and Prorocentrum minimum on early life history stages of the eastern oyster (Crassostrea virginica)

The bloom-forming dinoflagellates Prorocentrum minimum and Karlodinium veneficum can have detrimental effects on some marine life, including shellfish, but little is known about their effects on early life history stages of bivalves. In the Chesapeake Bay region, blooms of these dinoflagellates overlap with the spawning season of the eastern oyster, Crassostrea virginica. In laboratory experiments, we compared the effects of P. minimum and K. veneficum on the survival and development of embryos and larvae of the eastern oyster. At 10⁴ cells ml⁻¹, P. minimum did not have a negative effect on embryos and larvae in 2-day exposures. The yield of D-hinge larvae was equal to or greater than in control treatments. At 2 × 10⁴ cells ml⁻¹ (approximately equal biomass to the P. minimum treatment) K. veneficum caused significant mortality to oyster embryos within 1 day and almost no embryos developed into D-hinge larvae. This effect was not alleviated by the provision of an alternate food source (Isochrysis sp.). Significant mortality was observed when larvae were exposed to K. veneficum at concentrations of 10⁴ cells ml⁻¹ (approximately 5 ng ml⁻¹ of karlotoxin). The K. veneficum cultures used in these experiments were relatively low in toxin content, more toxic strains could be expected to cause mortality at lower cell concentrations. Survival and maturation of embryos and larvae may be reduced when spawns of the eastern oyster coincide with high bloom densities of K. veneficum.     

Terrestrial life of the amphibious fish Mnierpes macrocephalus

The physiological adaptations for terrestrial life of the rockskipper Mnierpes macrocephalus (Günther) have been examined and compared with other amphibious species. The rockskipper lives on steep rocky shores in the eastern tropical Pacific Ocean, and makes brief terrestrial excursions for orientation, feeding, and to avoid wave action and predators. The rockskipper spends a large portion of its time, and can live perfectly well, submerged in water. The duration of this species' daytime terrestrial sojourns rarely exceeds 30 min. While on land, the rockskipper respires aerially, through its gills and skin, at the same rate it normally respires in water. When exposed to sunlight without water, the rockskipper's body temperature increases rapidly. A small amount of water, however, enables the rockskipper to remain cool, apparently by dissipating absorbed heat through evaporative cooling of the gills and skin. Evaporative water-loss from respiratory surfaces is a major factor limiting terrestrial excursions of the rockskipper.     

Comparison of herbivory in the closely-related marine fish genera Girella and Kyphosus

  Specimens of the nominally herbivorous, closely-related, fish genera Girella and Kyphosus were collected from Australian waters in 1994 and 1995. The diet of three Girella species (G. cyanea, G. elevata, and G. tricuspidata) consisted mainly of chlorophytes and rhodophytes, with an animal component of␣15.9 ± 4.2% in G. tricuspidata. The diet of four species of Kyphosus (K. bigibbus, K. cinerascens, K. sydneyanus, and K. vaigiensis) included phaeophytes, chlorophytes and rhodophytes, and almost no animal material. Concentration of total short-chain fatty acids in the posterior intestine was <11.4 mM in the Girella spp. and >39.2 mM in the Kyphosus spp. These results suggest that microbial fermentation plays a role in algal digestion in Kyphosus spp., but not in Girella spp. Girellids and kyphosids appear to function quite differently as herbivores. Girellids should be considered as omnivores that complement readily-available energy from algae with protein from invertebrates. Kyphosids appear to be strict herbivores that can derive adequate nutrition from algae poor in easily assimilable energy, through microbial fermentation in the hindgut. Received: 8 July 1996 / Accepted: 2 August 1996     

Density dependence in social behaviour: home range overlap and density interacts to affect conspecific encounter rates in a gregarious ungulate

Sociality is poorly understood in the context of population processes. We used wild, female elk (Cervus canadensis) equipped with proximity-logging radio collars (n?=?62) from Manitoba, Canada (2007–2009), to test for modifying effects of population density (two areas: 0.42 and 0.22 animals/km²) on the relationship between two measures of sociality. This included the rate at which collared individuals encountered one another per year (encounters logged as animals ranging to within 1.4 m of each other) and the extent to which animals overlapped in annual home range (proportion of shared minimum convex polygon ranges). Overlap was significantly greater in the high density area compared to that of the low, but not if we only considered individuals that directly encountered each other, implying that familiar individuals will maintain a constant degree of range overlap regardless of density. Encounter rate was nonlinearly related to home range overlap. This relationship was also density-dependent, exhibiting negative density dependence at high proportions of overlap, primarily in the high density subpopulation. Sociality, as defined by two interacting measures of behaviour—encounter rate and home range overlap—exhibits a complex nonlinear relationship; we discuss the implications of these results as they pertain to sociobiology, resource competition, and pathogen transmission.     

Role of cortisol on condition factor in the female freshwater fish, Notopterus notopterus during four reproductive phases

The condition of the female fish, Notopterus notopterus was studied during four phases of the reproductive cycle in control and after cortisol hormone treatment. The condition of the fish including condition factor (K) and somatic condition factor (Ks) was determined based on weight of the body, length of the fish and gonad weight. In control fish the condition of the fish improve during prespawning phase compared to other phases. The hormone cortisol level estimated by radioimmunoassay (RIA) technique was found to be increased in preparatory and decreased during prespawning phase, may be because of the hormone involvement in metabolic activity, vitellogenesis. In cortisol treatment, the condition of the fish decrease during prespawning phase, compared to other phase may be because of extra expenditure of energy for progressing reproductive activity such as vitellogenesis.