Closing the larval loop: linking larval ecology to the population dynamics of marine benthic invertebrates

The majority of marine benthic invertebrates exhibit a complex life cycle that includes separate planktonic larval, and bottom-dwelling juvenile and adult phases. To understand and predict changes in the spatial and temporal distributions, abundances, population growth rate, and population structure of a species with such a complex life cycle, it is necessary to understand the relative importance of the physical, chemical and biological properties and processes that affect individuals within both the planktonic and benthic phases. To accomplish this goal, it is necessary to study both phases within a common, quantitative framework defined in terms of some common currency. This can be done efficiently through construction and evaluation of a population dynamics model that describes the complete life cycle.

Two forms that such a model might assume are reviewed: a stage-based, population matrix model, and a model that specifies discrete stages of the population, on the bottom and in the water column, in terms of simultaneous differential equations that may be solved in both space and time. Terms to be incorporated in each type of model can be formulated to describe the critical properties and processes that can affect populations within each stage of the life cycle. For both types of model it is shown how this might be accomplished using an idealized balanomorph barnacle as an example species. The critical properties and processes that affect the planktonic and benthic phases are reviewed. For larvae, these include benthic adult fecundity and fertilization success, growth and larval stage duration, mortality, larval behavior, dispersal by currents and turbulence, and larval settlement. It is possible to predict or estimate empirically all of the key terms that should be built into the larval and benthic components of the model. Thus, the challenge of formulating and evaluating a full life cycle model is achievable. Development and evaluation of such a model will be challenging because of the diverse processes which must be considered, and because of the disparities in the spatial and temporal scales appropriate to the benthic and planktonic larval phases. In evaluating model predictions it is critical that sampling schemes be matched to the spatial and temporal scales of model resolution.     

Individual-based modelling of recruitment variability and biomass production of bay anchovy in mid-Chesapeake Bay

Production of bay anchovy Anchoa mitchilli is highest in the larval and juvenile stages. The interplay between vital rates, stage durations, prey resources, and anchovy abundance ultimately determines the relative magnitude of recruitment (which in the model varies by about three-fold) and of stage-specific production. Changes in adult seasonal spawning patterns that increase the number of larval survivors result in only a slight increase in overall production due to density-dependent decreases in growth rates of later life stages. Bay anchovy in the mid-Chesapeake Bay may reach a compensatory threshold during late summer-autumn as fish growth is affected by competition for food resources. Density dependence in the population is evident in the relationships between spawner-recruit, size-recruit, and production of larval or juvenile to young-of-the-year life stages. Density-dependent growth acts differentially upon the early life stage that exceeds the compensatory threshold in any given year, due either to environmental variability or population size, or both. This could explain partially the relatively low recruitment variability observed for this anchovy.     

Survival against the odds: ontogenetic changes in selective pressure mediate growth-mortality trade-offs in a marine fish

For organisms with complex life cycles, variation among individuals in traits associated with survival in one life-history stage can strongly affect the performance in subsequent stages with important repercussions on population dynamics. To identify which individual attributes are the most influential in determining patterns of survival in a cohort of reef fish, we compared the characteristics of Pomacentrus amboinensis surviving early juvenile stages on the reef with those of the cohort from which they originated. Individuals were collected at hatching, the end of the planktonic phase, and two, three, four, six and eight weeks post-settlement. Information stored in the otoliths of individual fish revealed strong carry-over effects of larval condition at hatching on juvenile survival, weeks after settlement (i.e. smaller-is-better). Among the traits examined, planktonic growth history was, by far, the most influential and long-lasting trait associated with juvenile persistence in reef habitats. However, otolith increments suggested that larval growth rate may not be maintained during early juvenile life, when selective mortality swiftly reverses its direction. These changes in selective pressure may mediate growth-mortality trade-offs between predation and starvation risks during early juvenile life. Ontogenetic changes in the shape of selectivity may be a mechanism maintaining phenotypic variation in growth rate and size within a population.     

The effect of skewed distributions of vital statistics on growth of age-structured populations

Given that the variance of vital statistics can influence population projections, it seems reasonable that positive skew as observed for the distribution of larval survivorship of spruce budworms might also have a significant effect on stochastic projections of population growth. Simulations of population growth, using variable survivorship for a single age class, demonstrate that shape of the distribution of survivorship influences the outcome of stochastic population growth, and therefore is important for evolutionary and ecological theory. Unfortunately, empirical distributions of survivorships or fecundities for single life history stages are rare in the current literature.     

Asynchronous pupation of univoltine insects as evolutionarily stable phenology

Evolutionarily stable seasonal timing of larval feeding stages is studied theoretically for univoltine insects. In the evolutionarily stable (or ESS) population, each individual maximizes its own lifetime reproductive success by choosing the hatching and pupation dates, given the resource availability curve with a peak in the middle of a year, a higher daily mortality in the feeding stages, and the growth rate decreasing with the larval biomass in the population. If growth rate is proportional to the body size, the population at the ESS is composed of a mixture of phenotypes differing in hatching and pupation, but pupation interval over which some popation occur every day is much longer than hatching interval. If growth rate increases with the body size at a speed slower than linearly, large sized larvae should pupate earlier than small ones.     

Seasonal plasticity in growth and development of the speckled wood butterfly, Pararge aegeria (Satyrinae)

Regulation of growth and development by photoperiod was studied in a population of the speckled wood butterfly, Purarge aegeria L. (Lepidoptera: Satyrinae), from southern Sweden. Individuals were reared in a range of photoperiodic regimes (9L. to 22L) and temperatures (13°C to 21° C). Plasticity was found for important life-history traits- generation time, growth rate and final weight and seasonal regulation of development in response to photoperiod was found to occur at two levels. Purarge aegeria hibernates as a third instar larva or in the pupal stage, cantering one of four major developmental pathways in response to photoperiod: (1) direct development in both the larval and pupal stages, (2) pupal winter diapause with or (3) without a preceding larval summer diapause, or (4) larval winter diapause. In addition to this high-level regulation of individual development, larval growth rate and pupal development rate also appear to be finally regulated by photoperiod within each major pathway. As photoperiods decreased from 22 h to 17 h at 17° C, growth rate among directly developing larvae increased progressively, as was the case for larva? developing according to a univoltine life cycle from 17 h to 14 h. At two photoperiods, 13 h and 16 h (corresponding to shifts between major pathways), both larval and pupal development were extremely variable with the fastest individuals developing directly and the slowest developing with a diapause. This indicates a gradual nature of diapause itself, suggesting that the two level may not he fundamentally different.     

An experimental study of population regulation in the salamander,Notophthalmus viridescens dorsalis (Urodela: Salamandridae)

Summary The roles of density-dependent larval survival and cannibalism of larvae as potential mechanisms of population regulation in the newt (Notophthalmus viridescens dorsalis) were evaluated in laboratory and field experiments. In laboratory containers, adults cannibalized larvae and large larvae cannibalized smaller larvae. In artificial ponds, larval survival did not depend on initial larval density. No cannibalism could be demonstrated in the complex environment, although the experiment was powerful enough to detect an ecologically relevant difference in survival. Adult growth was negatively correlated with the final biomass of larval newts, suggesting that the two life stages competed for resources. Larval growth rates were negatively correlated with final larval density, suggesting that larvae competed with each other. The proportion of larvae that became sexually mature at age 7 months (paedomorphs and adults that skipped the eft stage) varied inversely with larval density. Therefore, the potential regulatory mechanisms identified in this study are competition within and between life stages.     

Temperature influences selective mortality during the early life stages of a coral reef fish

For organisms with complex life cycles, processes occurring at the interface between life stages can disproportionately impact survival and population dynamics. Temperature is an important factor influencing growth in poikilotherms, and growth-related processes are frequently correlated with survival. We examined the influence of water temperature on growth-related early life history traits (ELHTs) and differential mortality during the transition from larval to early juvenile stage in sixteen monthly cohorts of bicolor damselfish Stegastes partitus, sampled on reefs of the upper Florida Keys, USA over 6 years. Otolith analysis of settlers and juveniles coupled with environmental data revealed that mean near-reef water temperature explained a significant proportion of variation in pelagic larval duration (PLD), early larval growth, size-at-settlement, and growth during early juvenile life. Among all cohorts, surviving juveniles were consistently larger at settlement, but grew more slowly during the first 6 d post-settlement. For the other ELHTs, selective mortality varied seasonally: during winter and spring months, survivors exhibited faster larval growth and shorter PLDs, whereas during warmer summer months, selection on PLD reversed and selection on larval growth became non-linear. Our results demonstrate that temperature not only shapes growth-related traits, but can also influence the direction and intensity of selective mortality.     

Life History of Bittacomorpha clavipes (Fabricius) (Diptera: Ptychopteridae) in an Ozark Spring,U.S.A.

A population of Bittacomorpha clavipes was studied in an Ozark spring from July 1987- August 1988. Measurement of larval head capsules (N=4,544) showed B. clavipes has four distinct larval instars. Seasonal distribution of immature and adult life stages suggests that this population has an asynchronous, non-seasonal, multivoltine life history with at least three cohorts. First instar larvae and pupae were collected during all months except December through March and December through February, respectively. Adults and larval instars II-IV were collected throughout the year. Larval recruitment generally paralleled adult flight periods, and the presence of adults during all months sampled indicates a correlation with ovipositional periods. The life history documented here for B. clavipes suggests that limiting factors for growth and reproduction are not seasonally dependent for this population.     

Carry-over effects of the larval environment on post-metamorphic performance in two hylid frogs

Life history theory and empirical studies suggest that large size or earlier metamorphosis are suitable proxies for increased lifetime fitness. Thus, across a gradient of larval habitat quality, individuals with similar phenotypes for these traits should exhibit similar post-metamorphic performance. Here we examine this paradigm by testing for differences in post-metamorphic growth and survival independent of metamorphic size in a temperate (spring peeper, Pseudacris crucifer) and tropical (red-eyed treefrog, Agalychnis callidryas) anuran reared under differing larval conditions. For spring peepers, increased food in the larval environment increased post-metamorphic growth efficiency more than predicted by metamorphic phenotype and led to increased mass. Similarly, red-eyed treefrogs reared at low larval density ended the experiment at a higher mass than predicted by metamorphic phenotype. These results show that larval environments can have delayed effects not captured by examining only metamorphic phenotype. These delayed effects for the larval environment link larval and juvenile life history stages and could be important in the population dynamics of organisms with complex life cycles.     

Not all management is equal: a comparison of methods to increase wood turtle population viability

Management generally targets the most tractable life stage to rescue declining populations; however, that stage may not have the largest influence on recovery. Freshwater turtles are declining globally and early stages are frequently targeted for management, although the effectiveness of these actions on population growth are relatively unknown because of incomplete demographic data. We estimated the hatchling yearly survival rate for a freshwater turtle in the field using in situ enclosures to collect missing demographic information. We used these data to develop demographic models to calculate growth rate for a hypothetical, declining population of wood turtles (Glyptemys insculpta) in Wisconsin, USA, 2014–2019. We modeled growth for populations across a range of scenarios from no management to combinations of nest protection and head-starting at varying levels of effort. Nest protection alone did not increase population growth rate, while head-starting alone increased population growth by 0.07, with the largest increase in growth rate, 0.11, resulting from combinations of both approaches. No combination of nest protection and head-starting, without an increase in adult survival rate from the observed 0.88 to ≥0.95, led to population stabilization or increase. Populations of freshwater turtles, like the wood turtle, will likely only recover with a multi-faceted approach that targets multiple life stages simultaneously.     

Growth and the expression of alternative life cycles in the salamander Ambystoma talpoideum (Caudata: Ambystomatidae)

Complex life cycles (CLCs) contain larval and adult phases that are morphologically and ecologically distinct. Simple life cycles (SLCs) have evolved from CLCs repeatedly in a wide variety of lineages but the processes that may underlie the transition have rarely been identified or investigated experimentally. We examined the influence of larval growth rate on the facultative expression of alternative life cycles (metamorphosis or maturation as gill-bearing adults [= paedomorphosis]) in the salamander Ambystoma talpoideum. We manipulated growth rates by altering the amount of food individuals received throughout larval development. The expression of alternative life cycles in A. talpoideum is influenced by growth via food levels, but the same growth rates at different points in the larval period elicit different responses. Individuals were more likely to metamorphose (i.e. express a CLC) when food levels and growth rates were high later in development and more likely to mature without metamorphosing (SLC) when growth rates were comparatively low during the same point in development. Growth rates at particular points in development, rather than overall larval growth rate, may be an important proximate factor in salamander life-cycle evolution.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 639–646.     

Life history traits of pest and non-pest populations in the phytophagous ladybird beetle,Epilachna niponica (coleoptera, coccinellidae)

Life history traits of the phytophagous ladybird beetle,Epilachna niponica were compared between a non-pest population feeding on wild thistle and a pest population feeding on cultivated solanaceous crops, mainly potato. The pest population had larger females, a higher population growth rate, a more continuous oviposition schedule, and a shorter developmental period in the immature stages, compared with the non-pest population. The two populations showed no clear differences in phenology from spring to autumn, egg mass size, hatchability, and larval survival rate. Significant differences were found in number of eggs laid per female during the first and second periods, and in the developmental period of the immature stages. These life history traits were influenced primarily by food plant. A higher fecundity and shorter immature period appear to be readily altered by the host shift from thistle to potato.     

密度对大树蛙蝌蚪生长发育和存活率的影响

种群密度效应主要表现在影响和调节种群的死亡率、发育速率、繁殖率以及扩散、迁移等反应种群数量动态的重要参数。分析密度对大树蛙蝌蚪生长发育和存活率的影响,有助于探究外界环境因子导致大树蛙种群密度迅速下降的作用机制。设置了15、20、25只/L和30只/L 4个密度组,测量大树蛙蝌蚪的尾长、体长、发育时间和存活率。结果表明,在15—30只/L范围内,密度升高显著降低了大树蛙蝌蚪尾长和体长的生长速率,减小其变态期的尾长和体长大小,其中与15只/L相比,20、25只/L和30只/L 3组蝌蚪的尾长分别降低了11.6%、11.8%和13.9%,体长分别降低了11.1%、9.5%和12.9%;随着密度的升高,大树蛙蝌蚪发育至跗蹠部伸长期和前肢伸出期的所需时间显著延长;大树蛙蝌蚪生长后期的存活率随密度升高显著降低,但密度对蝌蚪生长早期的存活率影响不显著。因此,密度升高可显著减小大树蛙蝌蚪的尾长和体长、延长发育时间和降低其生长后期的存活率,可能影响大树蛙蝌蚪变态后的适合度。     

Life tables for worker honeybees

Life tables for worker honeybees covering all life span, and those for adults, were prepared for three seasonal cohorts, June bees, July bees and wintering bees. Survivorship curves for June and July bees show a convex type being exceptional for insects, with relatively high mortality at egg and feeding larval stages and at later adult stage after most bees became potential foragers. Adult longevity greatly lengthens in Winteriing bees and survivorship curve drops approximately with the same rate. A remarkable similarity of survivorship curves for men and honeybees was demonstrated, apparently due to highly developed social care in both. Some comments were given on mortality factors. The importance of life tables for population researches was shown by applying our result to the population growth curve made byBodenheimer , based upon the data byNolan . At the asymptote of the uncorrected curve, the ratio of total population estimated by uncorrected curve to that by corrected curve reaches about 3∶2.     

幼虫密度对甜菜夜蛾生长发育与繁殖的影响

为了研究甜菜夜蛾Spodoptera exigua(Hübner)幼虫的密度对其发育及繁殖的影响,本实验观察了5种幼虫密度下（1,5,10,20,30头/瓶）,幼虫发育和成虫繁殖情况。结果表明：幼虫和蛹历期、存活率和蛹重均差异显著。幼虫和蛹历期均以20头/瓶的最短,1头/瓶的最长,其余随幼虫密度增加而延长;幼虫至蛹存活率以10头/瓶的最高,其余随幼虫密度增加而降低;1头/瓶的蛹最重,显著高于其他密度的,其余随幼虫密度增加而下降。尽管密度间成虫羽化率和产卵前期均无显著差异,但成虫产卵量、寿命和畸形率差异显著。1头/瓶的产卵量最多,其次为10头/瓶的,其余随幼虫密度增加而减少,30头/瓶的产卵量显著少于其他密度的; 密度在1～20头/瓶范围内,雌蛾寿命均较短,显著短于30头/瓶的,而雄蛾寿命以5头/瓶的最短,显著短于其他密度的（10头/瓶除外）,10头/瓶的次之,其余密度间差异不显著;不同幼虫密度下羽化的成虫畸形率差异显著,10头/瓶的最低,其余随幼虫密度增加而升高; 生命表结果表明甜菜夜蛾在10头/瓶下世代存活率和种群增长指数均最高,幼虫密度过低或过高均不利于种群增长;世代存活率（S）和种群增长指数（I）与幼虫密度之间的关系均呈抛物线关系：S =－0.2087x²+2.5694x+211.52 (R²=0.88),I=－0.0552x²+0.9166x+54.168 (R²=0.95)。结果提示幼虫密度影响甜菜夜蛾种群动态的重要生态因子之一。     

Maladaptive changes in multiple traits caused by fishing: impediments to population recovery

Some overharvested fish populations fail to recover even after considerable reductions in fishing pressure. The reasons are unclear but may involve genetic changes in life history traits that are detrimental to population growth when natural environmental factors prevail. We empirically modelled this process by subjecting populations of a harvested marine fish, the Atlantic silverside, to experimental size-biased fishing regimes over five generations and then measured correlated responses across multiple traits. Populations where large fish were selectively harvested (as in most fisheries) displayed substantial declines in fecundity, egg volume, larval size at hatch, larval viability, larval growth rates, food consumption rate and conversion efficiency, vertebral number, and willingness to forage. These genetically based changes in numerous traits generally reduce the capacity for population recovery.     

Utilization of larval and pupal detritus by Aedes aegypti and Aedes albopictus

The utilization of detritus sources by mosquito larvae during development may significantly affect adult life history traits and mosquito population growth. Many studies have shown invertebrate carcasses to be an important detritus source in larval habitats, but little is known regarding how invertebrate carcasses are utilized by mosquito larvae. We conducted two studies to investigate the rate of detritus consumption and its effect on larval development and life history traits. Overall, we found that Aedes aegypti and Aedes albopictus larvae rapidly consumed larval detritus, while pupal detritus was consumed at a significantly slower rate. We also found that the consumption of larval detritus significantly increased larval survivorship and decreased male development time but did not significantly influence female development time or pupal cephalothorax length for either sex. Our results suggest that the direct consumption of larval detritus can support the production of adults in larval habitats that lack allochthonous detritus inputs or where such organic inputs are insufficient. These studies indicate that different forms of invertebrate detritus are utilized in distinct ways by mosquito larvae, and therefore different forms of invertebrate detritus may have distinct effects on larval development and adult life history traits.     

Responses of the western spruce budworm to varying levels of nitrogen and terpenes

Summary An agar diet study using western spruce budworm populations from Idaho and New Mexico was carried out to determine the effects of varying concentrations of nitrogen, beta-pinene, and bornyl acetate on larval growth and survival. Increased availability of nitrogen resulted in increased larval growth rate and survival to the adult stage. Larval growth rates from the Idaho population were higher on the high-nitrogen diet than were growth rates from the New Mexico population. The high level of beta-pinene improved larval growth at the high-nitrogen concentration. Bornyl acetate significantly reduced larval growth at both the low and high levels of nitrogen although the effect was greatest with the high-nitrogen diet. High bornyl acetate concentrations also significantly reduced survival to the adult stage. At high-nitrogen levels, a high concentration of bornyl acetate reduced larval growth rates and adult survival to a level similar to that occurring at the low nitrogen and low bornyl acetate concentrations.