Seasonal variation in the functional response of a coral-reef piscivore alters the inverse density-dependent mortality of its prey

Density-dependent processes are critical for regulating species’ populations, and piscivory of coral-reef fishes is frequently density dependent. However, the mechanism driving this density-dependent mortality is poorly understood, but may be caused by changes in a predator’s feeding rate at different prey densities (its functional response). An aquarium experiment replicated in winter and summer examined the functional response after 22 and 47 h of Cephalopholis cruentata feeding on Halichoeres pictus. With the exception of summer data after 47 h (density-independent mortality), mortality was inversely density dependent across all prey densities and increased with higher summer temperatures. The absence of an asymptotic pattern of inverse density-dependent mortality was caused by type II (summer) or dome-shaped type IV (winter) functional responses, with the benefits of schooling likely to cause the low mortality rates at higher prey densities. Predators’ functional responses may underlie the inverse density-dependent mortality reported in field studies of aggregating fishes.     

Simulating density-dependent relationships in south Texas northern bobwhite populations

Density dependence influences northern bobwhite (Colinus virginianus) reproduction and overwinter mortality. However, the functional forms of these density-dependent relationships or the factors that influence them during the annual life cycle events of this bird are not clear. We used a systems analysis approach with a compartment model based on difference equations (Δt = 3 months) for bobwhites in South Texas to simulate population behavior using 16 different functional forms of density-dependent production and overwinter mortality. During the reproductive season, a weak linear density-dependent relationship resulted in the longest population persistence (up to 100.0 yr), whereas a reverse-sigmoid density-dependent relationship had the worst population persistence (2.5–3.5 yr). Regarding overwinter mortality, a sigmoid or weak linear density-dependent relationship and a weak linear or no density-dependent reproduction relationship had the longest population persistence (87.5–100.0 yr). Weak linear density-dependent reproduction with either sigmoid or weak linear overwinter mortality produced stable fall population trends. Our results indicated that density dependence may have a greater influence on overwinter survival of bobwhites than previously thought. Inclusion of density-dependent functional relationships that represent both density-dependent reproduction and overwinter mortality, were critical for our simulation model to function properly. Therefore, integrating density-dependent relationships for both reproductive and overwinter periods of the annual cycle of bobwhite life history events is essential for conducting realistic bobwhite population simulation analyses that can be used to test different management scenarios in an integrated and interdisciplinary manner. © 2012 The Wildlife Society.     

Population regulation by post-settlement mortality in two temperate reef fishes

 Input of individuals dispersing into open populations can be highly variable, yet the consequences of such variation for subsequent population densities are not well understood. I explored the influence of variable input (”supply”) on subsequent densities of juveniles and adults in open local populations of two temperate reef fishes, the bluebanded goby (Lythrypnus dalli) and the blackeye goby (Coryphopterus nicholsii). Variable recruitment was simulated by stocking a natural range of densities of young fishes on replicate patch reefs. Density and mortality of the stocked cohorts were followed over time, until the fishes reached maturity. Over the first day of the experiments, mortality of both species was significantly density-dependent; however, there was still a very strong relationship between density on day 1 and density on day 0 (i.e., simulated recruitment was still an excellent predictor of population density). At this point in the study, the main effects of density-dependent mortality were to reduce mean densities and variation about the mean. Over the period from the start of the experiments until the time when maturity was reached by each species (about 1 and 3 months for Lythrypnus and Coryphopterus, respectively), mortality was strongly density-dependent. Such strong density-dependent mortality virtually eliminated any linear relationship between adult density and ”recruit” density. However, for both species, the relationship between these two variables was well fit by an asymptotic curve, with the asymptotic density of adults equal to c. 3/m² for Coryphopterus, and c. 10/m² for Lythrypnus. Natural recruitment (via settlement of larvae) to the reefs over the period of the study (9 months) was above the asymptotic densities of adults for the two species, even though the study did not encompass the periods of peak annual recruitment of either species. This suggests that adult populations of these two gobies may often be limited, and regulated, by post-settlement processes, rather than by input of settlers. Other studies have shown that mortality of the two species is density-independent, or only weakly density-dependent, on reefs from which predators have been excluded. Hence, it appears that predators cause density-dependent mortality in these fishes. Received: 26 November 1996 / Accepted: 5 April 1997     

Population stability in relation to resource availability in an introduced population of an herbivorous lady beetle

Mechanisms responsible for population stability in relation to resource availability were studied in an introduced herbivorous lady beetle,Epilachna niponica. The introduced population was relatively constant over a seven-year study period. Egg density was related to the variation in host-plant abundance in different years, and was highly stabilized during the period from reproductive adult to egg stage. Two density-dependent processes were identified in the reproductive season: (1) density-dependent reduction in fecundity and (2) density-dependent increase in female mortality and/or emigration, all of which operated early in the season. As a result, temporal variability in cumulative egg density was greatly reduced by mid-May, by which time approximately40% of total eggs were laid. A field cage experiment demonstrated that egg-laying of individual females was largely limited by resource availability even at low levels of leaf herbivory. Since movement activity of ovipositing females increased in a density-dependent manner, inter-plant movement is more likely to cause density-dependent reduction in fecundity and female loss, due to enhanced energy expendiditure. The introduced population was less stable than the source population, probably because of decreased inter-plant movement of females and the unlikelihood of egg resorption, both of which contribute significantly to the temporal stability ofE. niponica population densities.     

Gradual increase in the number ofAsphondylia aucubae (Diptera,Cecidomyiidae) due to inversely density-dependent mortality processes

The population dynamics of the aucuba fruit midge,Asphondylia aucubae (Japanese name: Aokimitamabae), were studied for 3 yr mainly at a broad-leaved evergreen forest on Mt. Shiroyama in Kagoshima City. This species is univoltine and adults emerge in May. Eggs were laid randomly inside the immature fruit of the host plant,Aucuba japonica (Japanese name: Aoki). Healthy fruit fell by the following March, whereas galled fruit remained on the trees even after the emergence of adult midges. The survival rate from the egg to adult stage was estimated to be 24.2%, and the number of midges on five census trees increased gradually during the census period. Two important mortality factors were recognized, i.e., fall of incompletely galled fruit and damage of the galled fruit by feeding of caterpillars of a moth. Some unknown factors were also found to be important, operating in an inversely density-dependent manner. The gradual increase ofA. aucubae is considered to be caused by such inversely density-dependent mortality processes.     

Density-dependent effects in a parasitic nematode,Elaphostrongylus rangiferi,in the snnail intermediate host

Summary Density-dependent effects in Elaphostrongylus rangiferi, a parasitic nematode in the CNS and muscular system of reindeer, were studied in a laboratory population of the snail intermediate host, Arianta arbustorum. The rates in parasite growth, development and mortality were all affected by parasite density. The effects on growth and development were, however, much more marked, than the effect on mortality.All density-dependent rates were intensified by decreasing snail size, and by snail starvation. The snail host showed marked tissue reactions against infection, and the intensity of these reactions increased with increasing parasite density. The mechanism behind the observed density-dependent rates is discussed, and is tentatively concluded to be competition for nutritive substances in the host tissue.The importance of a density-dependent developmental rate in natural populations of this parasite is discussed, and it is hypothesized that this effect may counteract the strong temperature-dependent developmental rate of E. rangiferi In a more general context it is pointed out that density-dependent developmental rates, although common amongst animal populations, has been neglected in models of population dynamics. Developmental rates are usually represented by a constant time lag in such models, but should be treated as a density-dependent variable.     

Density dependence: Applications in wildlife management

Knowledge of density-dependent processes is regarded as important for making decisions on the management of wildlife populations. Using published data on ungulates and upland game birds, we discuss density-dependent effects on population growth, harvest management under the logistic model, and management to increase or decrease survival and production. Empirical data show density-dependent growth for white-tailed deer (Odocoileus virginianus), reindeer (Rangifer tarandus), ring-necked pheasants (Phasianus colchicus), and northern bobwhites (Colinus virginianus), although the logistic model provided, at best, an approximation of growth. Managing harvest according to logistic theory is rare for ungulates and upland game; we suspect this owes to scarce data on population growth and complexity in density-dependent processes. Under density dependence, managing to increase production or survival may be self-defeating because an increase in 1 demographic variable entails a decrease in the other for sustaining populations (λ = 1). The problem can be addressed by providing space for population growth (λ > 1), at least until growth re-establishes the density-dependent response (λ = 1). © 2012 The Wildlife Society.     

Predators reverse the direction of density dependence for juvenile salmon mortality

The effect of predators on prey populations depends on how predator-caused mortality changes with prey population density. Predators can enforce density-dependent prey mortality and contribute to population stability, but only if they have a positive numerical or behavioral response to increased prey density. Otherwise, predator saturation can result in inversely density-dependent mortality, destabilizing prey populations and increasing extinction risk. Juvenile salmon and trout provide some of the clearest empirical examples of density-dependent mortality in animal populations. However, although juvenile salmon are very vulnerable to predators, the demographic effects of predators on juvenile salmon are unknown. We tested the interactive effects of predators and population density on the mortality of juvenile Atlantic salmon (Salmo salar) using controlled releases of salmon in natural streams. We introduced newly hatched juvenile salmon at three population density treatments in six study streams, half of which contained slimy sculpin (Cottus cognatus), a common generalist predator (18 release sites in total, repeated over two summers). Sculpin reversed the direction of density dependence for juvenile salmon mortality. Salmon mortality was density dependent in streams with no sculpin, but inversely density dependent in streams where sculpin were abundant. Such predator-mediated inverse density dependence is especially problematic for prey populations suppressed by other factors, thereby presenting a fundamental challenge to persistence of rare populations and restoration of extirpated populations.     

Density-related mortality of the flour beetle,Tribolium confusum duval

Life tables were constructed to assess the relative importance of some factors causing mortality of Tribolium confusum and to gauge their response to increasing population density. Observations focussed on three population densities (100, 400 and 800 individual/8 g medium) from the egg to the adult stage. The medium was not renewed in order to maximize predatory interactions. Generation mortality at densities 100, 400 and 800 was 42%, 50% and 74% respectively, i. e. density-dependent. Mortality in the first 10 days was also density-dependent reaching a maximum of 27% at density 800: predation by small larvae on eggs seemed the principal causative factor. The overall pattern of larval mortality was density-independent. Data on the mortality of pupae and callows were ultimately consistent with an inversely density-dependent pattern. Apparently, only mortality occurring within the first 10 days was capable of population regulation.     

Density dependence and the stabilization of animal numbers

Summary The claim of Varley and Gradwell that the highly density-dependent pupal predation (k ₅) in the population of the winter moth in Wytham Wood, England would keep density within limits (regulate) is compared in this paper with the density limits in the null model: pupal predation causes the same mean generation mortality (35%) as in the field, but is not density-dependent, i.e. either constant or randomly fluctuating between years according to the actual frequency distribution, ceteris paribus. According to this null model the winter moth would have fluctuated between narrower limits than in the field; the claim of Varley & Gradwell must thus be rejected. It is more generally concluded that a regulating factor should be the key factor, but this is not a sufficient condition. It should also prevent in some way the low throughs in the time series that usually accompany the operation of a dominating density-dependent mortality factor.Communication No. 264 of the Biological Station, Wijster     

What makes a species common? No evidence of density-dependent recruitment or mortality of the sea urchin Diadema antillarum after the 1983–1984 mass mortality

A potential consequence of individuals compensating for density-dependent processes is that rare or infrequent events can produce profound and long-term shifts in species abundance. In 1983–1984 a mass mortality event reduced the numbers of the abundant sea urchin Diadema antillarum by 95–99 % throughout the Caribbean and western Atlantic. Following this event, the abundance of macroalgae increased and the few surviving D. antillarum responded by increasing in body size and fecundity. These initial observations suggested that populations of D. antillarum could recover rapidly following release from food limitation. In contrast, published studies of field manipulations indicate that this species had traits making it resistant to density-dependent effects on offspring production and adult mortality; this evidence raises the possibility that density-independent processes might keep populations at a diminished level. Decadal-scale (1983–2011) monitoring of recruitment, mortality, population density and size structure of D. antillarum from St John, US Virgin Islands, indicates that population density has remained relatively stable and more than an order of magnitude lower than that before the mortality event of 1983–1984. We detected no evidence of density-dependent mortality or recruitment since this mortality event. In this location, model estimates of equilibrium population density, assuming density-independent processes and based on parameters generated over the first decade following the mortality event, accurately predict the low population density 20 years later (2011). We find no evidence to support the notion that this historically dominant species will rebound from this temporally brief, but spatially widespread, perturbation.     

Age-specific, density-dependent and environment-based mortality of a short-lived perennial herb

Density-independent and density-dependent processes affect plant mortality. Although less well understood, age-specific mortality can also play an important role in plant mortality. The goal of this study was to analyse several factors accounting for mortality in the Mediterranean short-lived perennial herb Lobularia maritima. We followed three cohorts of plants (from emergence to death) during 4 years in field conditions. We collected data on plant mortality of the effect of biotic agents (moth larvae and mycoplasma-like organisms, MLOs) and environmental variables. We also estimated density-dependent relationships affecting the fate of seedlings and adults. Results show that cohorts differed in their survival curves and ageing significantly increased mortality risk. Seedling mortality was density-dependent whereas adult mortality was not affected by density. MLO infection led to higher plant mortality whereas moth larvae attack did not affect plant mortality. In general, seedlings and adult plants experienced the highest mortality events in summer. We found, however, weak relationships between weather records and plant mortality. Age and size structures were not correlated. Overall, this study provides a comprehensive review of age-specific, density-dependent and density-independent factors that account for mortality of L. maritima plants throughout their life cycle in field conditions, highlighting the fact that age is an important factor in determining plant population dynamics.     

Density-dependent processes in the life history of fishes: evidence from laboratory populations of zebrafish Danio rerio

Population regulation is fundamental to the long-term persistence of populations and their responses to harvesting, habitat modification, and exposure to toxic chemicals. In fish and other organisms with complex life histories, regulation may involve density dependence in different life-stages and vital rates. We studied density dependence in body growth and mortality through the life-cycle of laboratory populations of zebrafish Danio rerio. When feed input was held constant at population-level (leading to resource limitation), body growth was strongly density-dependent in the late juvenile and adult phases of the life-cycle. Density dependence in mortality was strong during the early juvenile phase but declined thereafter and virtually ceased prior to maturation. Provision of feed in proportion to individual requirements (easing resource limitation) removed density dependence in growth and substantially reduced density dependence in mortality, thus indicating that 'bottom-up' effects act on growth as well as mortality, but most strongly on growth. Both growth and mortality played an important role in population regulation, with density-dependent growth having the greater impact on population biomass while mortality had the greatest impact on numbers. We demonstrate a clear ontogenic pattern of change in density-dependent processes within populations of a very small (maximum length 5 mm) fish, maintained in constant homogeneous laboratory conditions. The patterns are consistent with those distilled from studies on wild fish populations, indicating the presence of broad ontogenic patterns in density-dependent processes that are invariant to maximum body size and hold in homogeneous laboratory, as well as complex natural environments.     

Temporal changes in the strength of density-dependent mortality and growth in intertidal barnacles

1. In demographically open marine systems, the extent to which density-dependent processes in the benthic adult phase are required for population persistence is unclear. At one extreme, represented by the recruitment limitation hypothesis, larval supply may be insufficient for the total population size to reach a carrying capacity and density-independent mortality predominates. At the opposite extreme, populations are saturated and density-dependent mortality is sufficiently strong to reshape patterns established at settlement. 2. We examined temporal variation in the way density-independent and density-dependent mortality interact in a typical sessile marine benthic invertebrate, the acorn barnacle Semibalanus balanoides (L.), over a 2-year period. 3. Recruitment was manipulated at two high recruitment sites in north Wales, UK to produce recruit densities covering the range naturally found in this species. Following manipulation, fixed quadrats were monitored using digital photography and temporal changes in mortality and growth rate were examined. 4. Over a 2-year period there was a clear, spatially consistent, over-compensatory relationship between the density of recruits and adult abundance indicating strong density-dependent mortality. The strength of density dependence intensified with increasing recruitment. 5. Density-dependent mortality did not operate consistently over the study period. It only operated in the early part of the benthic phase, but the pattern of adult abundance generated was maintained throughout the whole 2-year period. Thus, early life-history processes dictated adult population abundance and dynamics. 6. Examination of the natural recruitment regime in the area of study indicated that both positive and negative effects of recruitment will occur over scales varying from kilometres to metres.     

Density-dependent resistance of the gypsy moth <Emphasis Type="Italic">Lymantria dispar</Emphasis> to its nucleopolyhedrovirus,and the consequences for population dynamics

The processes controlling disease resistance can strongly influence the population dynamics of insect outbreaks. Evidence that disease resistance is density-dependent is accumulating, but the exact form of this relationship is highly variable from species to species. It has been hypothesized that insects experiencing high population densities might allocate more energy to disease resistance than those at lower densities, because they are more likely to encounter density-dependent pathogens. In contrast, the increased stress of high-density conditions might leave insects more vulnerable to disease. Both scenarios have been reported for various outbreak Lepidoptera in the literature. We tested the relationship between larval density and disease resistance with the gypsy moth (Lymantria dispar) and one of its most important density-dependent mortality factors, the nucleopolyhedrovirus (NPV) LdMNPV, in a series of bioassays. Larvae were reared in groups at different densities, fed the virus individually, and then reared individually to evaluate response to infection. In this system, resistance to the virus decreased with increasing larval density. Similarly, time to death was faster at high densities than at lower densities. Implications of density–resistance relationships for insect–pathogen population dynamics were explored in a mathematical model. In general, an inverse relationship between rearing density and disease resistance has a stabilizing effect on population dynamics.     

Parasitism and predation as agents of mortality of winter moth populations in neglected apple orchards in Nova Scotia

Abstract.

• 1 This study compared the roles of pupal mortality and parasitism in winter moth (Operophtera brumata) population dynamics in Nova Scotian apple orchards and assessed the importance of beetles as pupal predators.
• 2 The component of pupal mortality termed predation accounted for greater stage-specific mortality of winter moth than parasitism by Cyzenis albicans in four neglected orchards.
• 3 Parasitism by Cyzenis albicans was not spatially density-dependent in any orchard, whereas the predation component of pupal mortality was spatially density-dependent in the two orchards most densely populated by winter moth.
• 4 Field experiments indicated that over 60% of pupal predation may be attributed to beetles, and that about 46% of pupal predation occurred within 4 weeks after pupal drop.
• 5 Mortality of introduced populations of winter moth in Nova Scotia resembles that of native populations in England where density-dependent predation regulates the winter moth population and reduces the parasitoid population to minimal levels. The situation in Nova Scotia appears to have changed appreciably since the establishment of parasitoids into the system in the 1950s.

     

The distribution of risk and density-dependent mortality in the galls of Eurosta solidaginis, the goldenrod gall fly

ABSTRACT.

• 1 This paper explores the net effect of a suite of mortality factors on a sedentary prey, the larvae of the goldenrod gall fly, Eurosta solidaginis Fitch (Diptera: Tephritidae).
• 2 Mortality is caused by unknown factors early in larval development, two species of parasitoid wasp (Hymenoptera: Eurytomidae), an inquiline beetle larva (Coleoptera: Mordellidae), and during the winter months downy woodpeckers Picoides pubescens (L.).
• 3 Distribution of mortality among galls relative to prey (gall) distribution was measured and discussed with respect to the distribution of relative risk of predation.
• 4 Galls are by and large contagiously distributed among quadrats, and mortality is distributed in a comparable pattern to that of galls.
• 5 The pattern of mortality on Eurosta larvae is neither density-dependent nor aggregated independently of gall distribution. Persistence in the system is probably a result of a combination of other factors such as adult mortality and early larval death which may have intergenerational density-dependent effects, and the linkage of locally unstable sub-populations via migration.

     

Population regulation in trichostrongylids of ruminants

Since regulation of population size requires the existence of one or more density-dependent processes affecting parasite numbers, the literature was examined for evidence of density-dependence in establishment, reproduction and mortality of trichostrongylids of ruminants. Because of differences between the two environments required for completion of the trichostrongylid life-cycle, this evidence was sought in processes occurring within the host, rather than in dung-pats or on pasture. Evidence was found of at least one density-dependent population process in several economically important species of the genera Haemonchus, Ostertagia and Trichostrongylus, although further observations on populations derived from continuous infection at several rates of larval intake are required to substantiate much of this evidence. It is suggested that populations of trichostrongylids of ruminants are regulated at the level of the suprapopulation by density-dependent constraints on egg production of constituent infrepopulations; rates of larval intake may play a central role in coordinating regulation within each ecosystem.     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

Biology ofhyphantria cunea drury (lepidoptera : Arctiidae) in Japan. IX. population dynamics

Fluctuations in population density of Hyphantria cunea in Japan are characterized by a gradation-like pattern. Analysis of the life table data taken from two stations during eight successive generations showed that (1) mortality during egg and early larval stages was density-independent, (2) mortality during later larval stages was inversely density-dependent, and (3) mortality during prepupal and pupal stages was density-independent. Thus the overall mortality process from egg to adults eclosion was inversely density-dependent. The inverse density-dependence in mortality process during later larval instars was mainly attributed to the ‘escape’ (VOÛTE, 1946) of H. cunea populations from the predation pressure of polyphagous predators such as birds and Politses wasps. This inverse density-dependence was considered to be a cause of gradation-like fluctuation. Field collection of egg-masses showed that the mean number of eggs per egg-mass, which was believed to be a good representation of mean fecundity, varied from 425 to 1050 during 4 years. Density-dependent reduction in the mean number of eggs per egg-mass was demonstrated, and this reduction was a factor regulating the population density. Assuming fixed sex ratio and survival rate of adults, a preliminary population model was constructed. The number of eggs laid in the survey station could be predicted well by the model based on the number of eggs laid in the previous generation, in 9 out of 13 cases. An attempt to apply a model of the same type to mimic the fluctuation of abundance (peak number of larval colonies per tree) on road-side trees suggested another density-dependent process, that is, insecticide application by man. Discussion was also presented on the causes responsible for the turn of population trend from decreasing to increasing in the 1st generation of 1968.