Modulation of predator-prey interactions by the Allee effect

An Allee effect arising from density-dependent mating success can have significant impacts at the ecosystem level when considered in the context of predator-prey interactions. These are captured by a mathematical model for the exchange of biomass between a structured predator population (continuous weight distribution) and a resource. Because the predator’s mating success affects the amount of resources required for the production of offsprings and their future growth into mature organisms, it influences the flux of biomass between trophic levels. Under simple assumptions, the equations can be reduced to an equivalent unstructured predator-prey model in which the Allee effect modulates the predation rate: the mating probability multiplies the rate of predator growth as well as the rate of resource depletion. Implications of the Allee effect for the bifurcation structure and equilibrium densities are examined. The model is compared to a modified version in which the Allee effect instead modulates the assimilation efficiency, hence the mating probability does not appear in the dynamical equation for the resource density. Both models exhibit qualitatively similar dynamics. However, compared to the model in which the Allee effect modulates predation, the model in which the Allee effect modulates assimilation efficiency predicts (i) unrealistically inefficient resource assimilation when predator density is low, (ii) a higher risk of catastrophic extinction resulting from a change in the parameter controlling the strength of the Allee effect, and (iii) no possibility of an increase in population size when the density dependence is enhanced.     

Invading parasitoids suffer no Allee effect: a manipulative field experiment

One frequent explanation for the failure of biological invasions is the Allee effect: due to positive density dependence, initially small invading populations may fail to establish and spread. Populations released for biological control are similar to fortuitous invading populations and may therefore suffer from Allee effects. However, unlike fortuitous invasions, biological control allows the experimental manipulation of initial population size and, thus, offers a unique opportunity to test for the occurrence of Allee effects. We manipulated the initial size of 45 populations of a parasitoid wasp introduced for the biological control of a phytophagous insect and followed the population dynamics of both parasitoids and hosts during three years. Our results suggest an absence of Allee effects but clear negative density dependence instead: (1) the probability of establishment after three years was not affected by initial population size; (2) net reproductive rate was highest at low parasitoid density and high host density; (3) the sex ratio, reflecting the proportion of virgin females, did not increase at low density, suggesting that low densities did not impede mate-finding; (4) the depression of host populations did not depend upon the number of parasitoids introduced. This is, to our knowledge, the first experimental test of the Allee effect in an invading parasitoid. It leads us to propose that a number of behavioral and life-history features of many parasitoids could protect them from Allee effects.     

Floater mortality within settlement areas can explain the Allee effect in breeding populations

The Allee effect (the positive relationship between population growth rate and population size) is a constraint of some animal populations at low numbers, which increases their likelihood of extinction because of a decrease in reproduction and/or survival. We were able to demonstrate that the Allee effect can be the result of a mortality increase affecting floaters (i.e. dispersing individuals able to enter as breeders in the reproductive population when a breeding territory or a potential mate – owner of a suitable breeding territory – becomes available). Previously, potential mechanisms underlying Allee effects were always related to the breeding portion of a population only. In contrast, our understanding of or solutions to population declines due to the Allee effects can reside elsewhere, away from breeding territories.     

Potential effect of phytoplankton colony breakage on the calculation of zooplankton filtration rates

Substantial net production of short chains of the diatom Skeletonema costatum occurred in grazing experiments conducted with Acartia hudsonica (=A. clausi). A general model was then constructed to evaluate the influence of colony breakage during grazing on the calculation of zooplankton filtration and ingestion rates. Breakage can lead to either over-or underestimation of these rates. The magnitude of error is related to breakage probability, particle size distribution, size of the largest colonies, length of the experiment, copepod concentration and actual filtration rates. Size distributions similar to those described in the literature as manifestations of a complex selection for size and particle abundance can be generated by the grazing model under conditions of a uniform filtration rate accompanied by colony breakage. Unless the rate of breakage exceeds the rate of removal and causes net particle production, the occurrence of breakage may go undetected.     

Experimental Demonstration of an Allee Effect in American Ginseng

Abstract: Harvesting of wild American ginseng (   Panax quinquefolius ) for the herbal trade has lowered natural population sizes. We tested for reproductive limitation due to small population size (a form of the Allee effect) by experimentally planting "natural" populations numbering 4, 16, and 64 using 4-year-old cultivated plants. Plant size traits and reproductive traits ( bud, flower, green fruit, and mature fruit) were recorded through the ensuing summer. Fruit production per flower and per plant increased in proportion to flowering population size (  p = 0.0063 and p = 0.0017, respectively), strongly suggesting that an Allee effect occurs in very small populations. The increase in fruit production was not explained by either plant or inflorescence size differences. Although population size-dependent pollination, through insufficient pollinator visitation rate or pollen transfer rate, seems the most likely cause of the observed effects, our limited observations of pollinators were not sufficient to demonstrate a change in pollination rates as a function of population size. Knowledge of the presence as well as the mechanism underlying this Allee effect may be especially useful for management and determination of minimum viable population size of the species in the wild.     

A curvilinear effect of height on reproductive success in human males

Human male height is associated with mate choice and intra-sexual competition, and therefore potentially with reproductive success. A literature review (n = 18) on the relationship between male height and reproductive success revealed a variety of relationships ranging from negative to curvilinear to positive. Some of the variation in results may stem from methodological issues, such as low power, including men in the sample who have not yet ended their reproductive career, or not controlling for important potential confounders (e.g. education and income). We investigated the associations between height, education, income and the number of surviving children in a large longitudinal sample of men (n = 3,578; Wisconsin Longitudinal Study), who likely had ended their reproductive careers (e.g. > 64 years). There was a curvilinear association between height and number of children, with men of average height attaining the highest reproductive success. This curvilinear relationship remained after controlling for education and income, which were associated with both reproductive success and height. Average height men also married at a younger age than shorter and taller men, and the effect of height diminished after controlling for this association. Thus, average height men partly achieved higher reproductive success by marrying at a younger age. On the basis of our literature review and our data, we conclude that men of average height most likely have higher reproductive success than either short or tall men.     

The effect of patch isolation on epifaunal colonization in two different seagrass ecosystems

The macrofaunal colonization of isolated habitats is affected by many factors, ranging from distance to the nearest source population to the dispersal mechanism of the species. We investigated the initial epifaunal colonization at two sites, one situated in the Northern Gulf of Mexico and the other in the Northern Baltic Sea. At each site, artificial seagrass units were placed at 10- and 20-m distances from a continuous seagrass meadow, as well as inside the meadow over a 5-day colonization time. With the exception of amphipods in Gulf of Mexico, patch isolation had a negative effect on colonization for the other faunal species, irrespective of the sites. This inverse colonization pattern of amphipods suggests that they are not equally sensitive to patch isolation in different regions. Our results indicate that increasing habitat isolation can have serious consequences for the community composition of seagrass epifauna. Furthermore, we emphasize the need for larger-scale latitudinal comparative studies.     

Small-scale horizontal distributions of zooplankton taxa

Small-scale (100 to 2 400 m) horizontal distributions of major taxonomic categories (class and order) of zooplankton were measured at a depth of 90 m with an opening-closing plankton net over a 3 d period in October 1978 in the California Current. Some zooplankton categories showed evidence of diurnal vertical migration, while others had long-period temporal changes in mean abundance. Variance-to-mean ratio for large copepods and euphausiids was higher at night than during the day, while the opposite was true for chaetognaths and pteropods. Within a given category, the variance-to-mean ratio generally increased with a category's abundance. Spatial abundance variations were characterized by trends (i.e., fluctuations larger than length of the net hauls) in some taxonomic categories. No consistent differences in scales of variability were found as a function of animal size or from day to night. Correlation analysis of taxonomic counts implied that significant biological interactions occurred. The proportions of counts of taxonomic groups showed no large changes over the time-space scales sampled. However, the proportions of biomass in taxonomic groups differed from day to night due to the large variability of euphausiids. Comparisons of wet weight biomass to taxonomic counts indicated that biomass was usually less variable than taxonomic counts.     

The consequences of being colonial: Allee effects in metapopulations of seabirds

Most seabirds live in large colonies. This fact signifies that there is an advantage in living and breeding together. Four explanations are put fore ward for this colonial behaviour, more birds have: (1) a reduced per capita predation of chicks in colonies, (2) a better anti-predator defence, (3) a more efficient foraging in temporally patchy environments and (4) sex ratios that are more likely to be close to one. These factors induce a strong Allee-type density-dependent relation, a positive relation between density and population growth rate at low density. Nevertheless, these Allee effects are generally ignored in seabird population studies. Therefore we study the consequences of introducing Allee-type density-dependent relations in a spatially explicit metapopulation model for the Common Tern (Sterna hirundo). Simulations show that Allee effects might be responsible for a 20-fold decline in the recolonization distances, causing patches and parts of metapopulations to effectively become more isolated. This leads to long recolonization times of empty breeding patches which consequently cause slower metapopulation expansion and recovery. Additionally, we show that the typical early warning signals, that show that a population is near its critical threshold induce by Allee effects, is less pronounced in colonies that are part of a metapopulation. Hence, we offer some simple equations to estimate critical densities and thresholds in a colony.     

The effect of ocean acidification on early algal colonization stages at natural CO2 vents

Marine algae exhibit different responses to ocean acidification, suggesting that a decrease in pH does not always favour marine photosynthetic organisms. In order to understand the effect of acidification on algal community development, early colonization stages were investigated using carbon dioxide vents around the Castello Aragonese (Ischia, Italy) as a natural laboratory. Settlement tiles were placed in zones with different pH (normal, medium and low), and species composition and coverage measured after 2, 3 and 4 months of deployment. The number of species decreased by 4 and 18 % at medium and low pH zones, respectively (P < 0.05). The structure of the algal assemblage differed between pH zones during the 4 months of the experiment, due to the addition and/or replacement of new species. This leads to a change in the succession of morphological forms as soft crustose algae replaced calcareous species, and turf species were dominant in cover; more complex thalli started to occur only at medium pH. These results support previous findings that ocean acidification will induce changes in benthic algal communities.     

重污染河道浮游动物种群的特点

苏州城市河道水情复杂,氮磷浓度较高,属于重污染的滞流水体.通过对试验区南园水系为期1a的调查研究表明:该河段共有浮游动物33种,其中原生动物4种,轮虫类24种,枝角类4种,桡足类1种.各河段浮游动物年平均密度为678 ind·L-1,生物量为56.028 mg·L-1.从种类数来看,轮虫类大于原生动物大于枝角类大于桡足类.从密度看,全年平均以轮虫最多,其次为原生动物,桡足类最少.从生物量看,以枝角类最高,桡足类次之,原生动物最少.从季节变化来看,春夏季较多,秋冬季较少.优势种为轮虫类和枝角类,这些种类极耐有机污染,耐低溶氧.在苏州古城区重污染河道中,浮游动物的总量与水体理化指标及浮游植物的数量等均没有线性相关性(差异显著,p<0.05),这可能与苏州城市河道污染较重有关.城市河道特殊的水域环境决定了浮游动物种群的形成、演变及其发展都有其特殊性.探讨了重污染的城市河道的浮游动物种类组成特点,提出了控制某些优势种群发展的设想,为改善河道水环境提供了理论依据.     

Global patterns of epipelagic gelatinous zooplankton biomass

There is concern that overfishing may lead to a proliferation of jellyfish through a process known as fishing down the food web. However, there has been no global synthesis of patterns of gelatinous zooplankton biomass (GZB), an important first step in determining any future trends. A meta-analysis of epipelagic-GZB patterns was undertaken, encompassing 58 locations on a global scale, and spanning the years 1967–2009. Epipelagic-GZB decreased strongly with increasing total water column depth (r ² = 0.543, p < 0.001, n = 58): in shallow (<50 m) coastal waters, epipelagic-GZB was typically 742× the levels in deep ocean (>2,000 m) sites. However, the ratio of GZB to primary productivity showed high values across a range of depths, i.e. this measure of the relative abundance of gelatinous zooplankton did not co-vary with depth.     

Metabolic characteristics of midwater zooplankton: Ammonia excretion,O:N rations,and the effect of starvation

The nature of protein catabolism in a wide range of species of midwater zooplankton was investigated. The weight-specific ammonia excretion rates (g NH₃–N g^–1 dry wt h^–1, y) decline exponentially with minimum depth of occurreece (MDO, x), y=163.4 x^{–0.479±0.212} (95%ci) (CI=confidence interval), when temperature is held constant. The change in ammonia excretion can be partially explained by the decrease in percent protein (%P) with MDO, %P=80.17 MDO^{–0.148±0.122 (95%ci)} The atomic O:N ratio of freshly caught zooplankters ranged from 9.1 to 91, with most measurements between 9 and 25. Detailed studies were carried out on the response of one of the species studied (Gnathophausia ingens) to starvation (28 d). After 14 d of starvation the average ammonia excretion rate declined by more than 75% to less than 1 g NH₃–N g^–1 wet wt h^–1, although the average oxygen consumption declined by only 13% within the first 7 d of starvation and then remained stable. This differential response of oxygen consumption and ammonia excretion to starvation resulted in an increase in the average O:N ratio of starved animals from an initial 33 to 165 after 21 d. The average O:N ratios of fed mysids remained below 38 during the experiment. G. ingens maintains a relatively uniform metabolic rate during starvation by relying more heavily on its large lipid stores than when being fed.     

富营养化水体中浮游动物对藻类的控制作用

采用室内受控生态系统进行实验.采用于桥水库原水接种后,人工设置营养盐质量浓度梯度,模拟富营养化程度不同的水体,以研究浮游动物的生长情况,及其控制藻类生长的作用.使用SPSS统计软件进行数据分析.结果显示:过高的营养盐浓度,可能会对轮虫的生长产生抑制作用,而甲壳类动物的总数量基本由营养盐质量浓度决定.在四个营养盐质量浓度梯度下,浮游动物与藻类之间均存在一定的显著或极显著相关关系,说明浮游动物均能够发挥一定的控藻作用.当营养盐ρ(TN)=3mg·L-1、ρ(TP)=0.02mg·L-1左右,浮游动物的控藻作用最明显.当营养盐质量浓度过高时,浮游动物的控藻作用受影响.     

Small-scale pattern of a California current zooplankton assemblage

The small-scale vertical (10s of meters) and horizontal (100s to 1000s of meters) distribution of zooplankton in the California Current near Guadalupe Island, Baja California, Mexico was studied. Vertical distributions were sampled using a vertically-towed Longhurst-Hardy Plankton Recorder (LHPR) which gave a sequence of samples, each integrated over about 5 m, from 250 m to the surface. Because of the sampling biases of the LHPR, details in vertical structure of less than 15 to 20 m were not considered. The pattern of the vertical tows and the variability of the integrated counts of species were used to infer horizontal distribution. Four series of 8 tows each were taken around noon and midnight over a 2-day period. Four of the 8 tows in each series were randomly positioned within 2000 m of a parachute drogue (first day) or a fixed geographic position (second day); four were replicate tows taken at the drogue or the fixed position. Sixty-seven taxonomic categories were counted. The replicate tows, separated by no more than a few hundred meters, gave more similar vertical profiles for species than did the random tows, with separations of 100s to 1000s of meters. The night replicate tows showed less variability in depth distribution than did either the night random or any of the day tows, leading to the hypothesis that the vertical distributions observed were generated by interactions of the organisms' diel behavior with internal waves. Variability of abundance estimates using the integrated counts was the same for both replicate and random tows, indicating that horizontal patches may be smaller than 100 m. No evidence was found for a day-night change in patch size, or for a consistent overlapping of patches of different species. Replicate tows gave more similar estimates of community structure (relative proportions of species) than did random tows. Overall day community structure was more similar between tows than night structure. Similarities in species' proportions of any random tow to the replicate tows or to other random tows of a series decreased with increasing distance between the tows being compared. This decrease was greater for the night samples, suggesting that community structure is more heterogeneous at night.This study is based in part on a dissertation submitted in partial fulfillment of the requirements for the Ph.D. degree at the University of California, San Diego, and was supported by National Science Foundation Grant GB 12413 and the Marine Life Research Program of the Scripps Institution of Oceanography.     

Nocturnal movements of phototactic zooplankton in shallow waters

Downwardly directed light-traps were used by night, over seagrass and coral sand to investigate the existence and significance of nocturnal peaks in phototactic zooplankton. Samples each half-hour from sunset to sunrise showed that small invertebrates, mostly copepods, were trapped in large numbers once or twice a night. Each period of high catches lasted only 1 to 2 h and was often associated with periods of failing natural light (evening twilight, moonset). This pattern and its variations are explained in terms of nocturnal vertical migration, in which plankton move towards light in response to decreasing light intensity, move away from light in response to increasing light intensity, and disperse at constant light intensity. Some morning peaks are unexplained.     

Nutrient regeneration by zooplankton during a red tide off Peru,with notes on biomass and species composition of zooplankton

During March and April 1976, a red tide, dominated by the dinoflagellate Gymnodinium splendens Lebour, developed in the vicinity of 15°06'S and 75°31'W off Peru. At the height of the bloom, the euphotic zone was 6 m deep and the chlorophyll a at the surface was 48 g l^-1. A daily collection of zooplankton at 09.00 hrs showed large fluctuations of biomass, from 0.2 to 3.84 g dry weight m^-2 in a water column of 120m. Copepodids and nauplii dominated the collections. During a period of reduced wind, the adult copepods were a mixture of the species characteristic of the coastal upwelling system and the neritic species associated with more northerly, tropical waters. Nitrogen regeneration by the zooplankton varied with the development of the bloom, the type of zooplankton dominating the experiment, and biomass fluctuations, but never accounted for more than 25% of the nitrogen uptake by phytoplankton.     

The effect of laboratory conditions on the extrapolation of experimental measurements to the ecology of marine zooplankton. IV. Changes in respiration and excretion rates of boreal zooplankton species maintained under fed and starved conditions

Herbivorous zooplankton species (Calanus plumchrus, Paracalanus parvus and Euphausia pacifica) and carnivorous species (Parathemisto pacifica and Pleurobrachia pileus) collected from Saanich Inlet, British Columbia, Canada, were maintained in the laboratory under fed and starved conditions. Respiration rate and excretion rates of ammonia and inorganic phosphate were measured successively on the same batch populations of each species in different feeding conditions. Respiration rate remained at a constant level or increased during the feeding experiment but decreased progressively in starved individuals. Herbivorous, but not carnivorous, species showed a rapid decrease in both excretion rates for the first few days of an experiment irrespective of feeding conditions. However, the general level of excretion rates of fed specimens was higher than that of starved ones. The O:N, N:P and O:P ratios were calculated from respiration, ammonia excretion and phosphate excretion and discussed in relation to metabolic substrates of animals during the experiment. A marked difference was shown in the O:N ratio between fed hervivores (>16) and fed carnivores (7 to 19), suggesting highly protein-oriented metabolism in the latter. One unknown factor causing variation in excretion rates is speculated to be the physiological stress on animals during sampling from the field. It is suggested that the laboratory measurement of realistic excretion rates of zooplankton is difficult owing to their large fluctuations, but this is not the case with respiration rate.     

In situ grazing rates of deep-sea benthic boundary-layer zooplankton

In situ grazing rates of mixed assemblages of deep-sea benthic boundary-layer zooplankton were measured in July 1983 at a site in the North Atlantic Ocean at 2 175 m depth using a short-term radioisotope-uptake method. Zooplankton were collected with an opening-closing net system from the bottom 1 m of the ocean and incubated in situ with mixed tritiated amino acids in special cod-end chambers. Incubations were terminated at depth by the addition of MS-222. Radioisotope uptake beyond that of dead controls was shown by both the zooplankton and particulate fractions. Grazing rates in the deep-sea experiments were surprisingly high, being comparable to wintertime Narragansett Bay zooplankton grazing rates determined in a separate series of laboratory experiments. These laboratory experiments also documented nonparticle-associated uptake of dissolved amino acids by Narragansett Bay zooplankton, but the importance of this in the deep sea is unknown. The deep-sea benthic boundary layer may be a region of elevated rates and activity because of its higher particulate concentrations, and our experiments may also have measured maximum rather than average rates.