A general formula for the sensitivity of population growth rate to changes in life history parameters.

This paper considers the sensitivity of population growth to small changes in birth, growth, survival, and migration probabilities for an arbitrary population classification (i.e., age, instar, size, developmental stage, age, and spatial location, etc.). The stage-specific life history parameters are expressed in a discrete-time system of linear difference equations, the dominant eigenvalue of which defines the population growth rate. The sensitivity of this eigenvalue to production of class i by class j individuals is shown to be proportional to the product of the reproductive value of stage i and the abundance of stage j in the stable stage distribution. This formula is readily computable, and several examples are presented. For the special case of age-structured populations, this formula reduces to those derived by Hamilton, Emlen, and Goodman.     

Early life history of the Mediterranean gorgonian Paramuricea clavata: implications for population dynamics

Abstract. Knowledge about early stages of marine sessile invertebrates dispersing by means of free-swimming propagules is fundamental toward understanding their population biology. In this study, we describe and quantify survivorship of early stages of the gorgonian Paramuricea clavata to contribute to the understanding of their implications on population dynamics of this emblematic species of the Mediterranean hard-bottom communities. Spawning was recorded in June for the 3 years studied (2001–2003). High levels of fertilization rate were observed during the main spawning in 2001 and 2002 (62–69%). This appears to be related to the surface brooding mode of development, synchronous spawning, and high fecundity of the species. The timing of development of the blastulae was ∼24 h and the planulae appeared after 48–72 h. In the laboratory, metamorphosis into polyp started between 8 and 25 d, but this timing may have been delayed by lack of unknown appropriate cues for settlement. The behavior of first developed planulae exhibited a marked negative phototaxis that may be a strategy to avoid competition with fast-growing algae on photophilous habitats. Despite the high fertilization rate in P. clavata , laboratory and field experiences indicate that survivorship through the planulae and polyp stages was very low, given that none of the settled polyps survived for >7 months in both experimental years. The low survival of the early stages may help explain the low recruitment rates observed in the field in addition to indicating major limitations for colonization of new areas and for persistence of the local population under disturbances.     

Determinate growth and modularity in a gorgonian octocoral

Growth rates of branches of colonies of the gorgonian Pseudopterogorgia elisabethae were monitored for 2 years on a reef at San Salvador Island, Bahamas. Images of 261 colonies were made at 6-month intervals and colony and branch growth analyzed. Branch growth rates differed between colonies and between the time intervals in which the measurements were made. Colonies developed a plumelike morphology through a pattern of branch origination and determinate growth in which branch growth rates were greatest at the time the branch originated and branches seldom grew beyond a length of 8 cm. A small number of branches had greater growth rates, did not stop growing, and were sites for the origination of subsequent "generations" of branches. The rate of branch origination decreased with each generation of branching, and branch growth rates were lower on larger colonies, leading to determinate colony growth. Although colonial invertebrates like P. elisabethae grow through the addition of polyps, branches behave as modules with determinate growth. Colony form and size is generated by the iterative addition of branches.     

Effects of temperature on life history variables in perch

Population structure, recruitment, mortality, life expectancy, age at maturity, gonad development, fecundity, net reproductive rate and energy storage were studied in a perch population exposed to thermal discharge. The experiment was conducted during 1978–1990 in an artificial enclosure at a nuclear power plant on the Baltic Sea coast. Abundance increased after the plant started operating, although the proportion of larger perch dropped to a low level. Recruitment improved, but adult mortality increased. This effect was accentuated as the fishes matured very early and at a very small size, reducing their ability to support a growing gonad in high winter temperatures. During the spawning period, condition dropped to very low levels accompanied by increased mortality. Surviving fish delayed the next spawning by one or more years and reduced their fecundity. Body condition thus improved in large perch at the end of the study period. Life-time fecundity was reduced, and reproductive performance shifted to youner ages.     

The dual role of sclerites in a gorgonian coral: Conflicting functions of support and defence

Sclerites in the gorgonian coral Briareum asbestinum perform the dual role of skeletal support against wave action and structural defence against predators. Local populations of B. asbestinum vary along gradients of decreasing water movement and decreasing predator abundance with increasing depth, such that sclerite length increases and sclerite density decreases with depth. Based on this pattern, I explored a possible trade-off between the sclerite composition that is most resistant to tearing versus most deterrent to predatory gastropods. Feeding assays revealed that artificial foods containing longer sclerites and those containing higher volume fractions of sclerites are less palatable to the gastropod Cyphoma gibbosum. However, real colonies appear constrained, in that they do not contain both long sclerites and high volume fractions at the same time. Given a choice among real colonies, snails prefer shallow-water colonies with shorter sclerites e ven though the sclerite volume fractions are high. Although least deterrent to snails, shallow-water colonies are 56% more resistant to tearing than their deep-water counterparts. Hence, variation in sclerite composition among local populations of B. asbestinum may be maintained by opposing selection for support versus defense.     

Reproductive styles and life history variables relative to exploitation and management ofSebastes stocks

Synopsis The characteristics of lightly and heavily exploited Pacific ocean perch,Sebastes alutus, stocks are evaluated relative to the predictions of life history theory. These long-lived species (50–100 year lifespan) show limited phenotypic plasticity and have little buffering against the effects of reduced lifespan. Reduced stock abundance has generated some compensatory increase in growth rate. Length at first maturity varies only slightly with increased growth rate, although age at maturity may decrease by 1–4 years. Grooth increases yield larger (15–20%) size at age and increased reproductive effort at younger ages, but lower size-specific fecundity for these faster-growing fish. This suggests an energy allocation protocol favouring growth over reproduction in these long-lived animals. Rockfishes have late recruitment to fisheries (ages 10–15), and the detection time for results of management actions is equally long. Their vulnerability to overfishing means that indices of population changes, more representative of fishing effects than the catch rate index presently used, are required. Reproductive value indices are shown to be extremely sensitive and continuous with population abundance changes. Their incorporation into monitoring programs would permit more timely evaluation of management actions. Management policies developed for shorter-lived species are shown to be inappropriate for rockfishes. The need for an improved match in the time frame of the species' life history, and that of management strategies, is stressed.     

Lifetime growth in wild meerkats: incorporating life history and environmental factors into a standard growth model

Lifetime records of changes in individual size or mass in wild animals are scarce and, as such, few studies have attempted to model variation in these traits across the lifespan or to assess the factors that affect them. However, quantifying lifetime growth is essential for understanding trade-offs between growth and other life history parameters, such as reproductive performance or survival. Here, we used model selection based on information theory to measure changes in body mass over the lifespan of wild meerkats, and compared the relative fits of several standard growth models (monomolecular, von Bertalanffy, Gompertz, logistic and Richards). We found that meerkats exhibit monomolecular growth, with the best model incorporating separate growth rates before and after nutritional independence, as well as effects of season and total rainfall in the previous nine months. Our study demonstrates how simple growth curves may be improved by considering life history and environmental factors, which may be particularly relevant when quantifying growth patterns in wild populations.     

Population dynamics and life history of a geographically restricted seahorse, Hippocampus whitei

The aim of this study was to collect data on population dynamics and life history for White's seahorse Hippocampus whitei, a geographically restricted species that is listed as data deficient under the IUCN Red List. Data from H. whitei populations were collected from two regions, Port Stephens (north) and Sydney Harbour (south) in New South Wales, Australia, covering most of the known range of H. whitei, from 2005 to 2010. Over 1000 individuals were tagged using fluorescent elastomer and on subsequent recaptures were re-measured for growth data that were used in a forced Gulland-Holt plot to develop growth parameters for use in a specialized von Bertalanffy growth-function model. Growth parameters for Port Stephens were: females L(∞) = 149·2 mm and K = 2·034 per year and males L(∞) = 147·9 mm and K = 2·520 per year compared with estimates from Sydney Harbour: females L(∞) = 139·8 mm and K = 1·285 per year and males L(∞) = 141·6 mm and K = 1·223 per year. Whilst there was no significant difference in growth between sexes for each region, H. whitei in Port Stephens grew significantly quicker and larger and matured and reproduced at a younger age than those from Sydney Harbour. The life span of H. whitei is at least 5 years in the wild with six individuals recorded reaching this age. Data collected on breeding pairs found that H. whitei displays life-long monogamy with three pairs observed remaining pair bonded over three consecutive breeding years. Baseline population densities were derived for two Port Stephens' sites (0·035 and 0·110 m(-2) ) and for Manly in Sydney Harbour (1·050 m(-2) ). Even though the life-history parameters of H. whitei suggest it may be reasonably resilient, precaution should be taken in its future management as a result of its limited geographical distribution and increasing pressures from anthropogenic sources on its habitats.     

Anthropogenic harvesting pressure and changes in life history: insights from a rocky intertidal limpet

The importance of large breeding individuals for maintaining the health of marine fish and invertebrate populations has long been recognized. Unfortunately, decades of human harvesting that preferentially remove larger individuals have led to drastic reductions in body sizes of many of these species. Such size-selective harvesting is particularly worrisome for sequentially hermaphroditic species where the larger size classes are composed primarily of one sex. Whether these species can maintain stable sex ratios under sustained harvesting pressure depends on the level of plasticity of their life-history traits. Here, we show that populations of a marine limpet (Lottia gigantea) can adjust a fundamental aspect of their life history (the timing of sex change) when subjected to size-selective harvesting. As predicted by theoretical models, individuals from harvested populations change sex at smaller sizes and grow at slower rates compared to individuals from protected populations. In addition, the relative size at which the change from male to female occurs remains constant (~0.75; size at sex change/maximum size) across populations, regardless of harvesting pressure. Our results show that population-level demographic and life-history data, in conjunction with existing theory, can be sufficient to predict the responses of sequential hermaphrodites to harvesting pressure. Furthermore, they suggest such species can potentially adapt to size-selective harvesting.     

Population regulation and the life history studies of LaMont Cole

Transformative changes marked the growth of mid-twentieth century American ecology. This included redirection of the scholarly focus of the discipline, especially on the role evolutionary theory and "levels of selection", and increased visibility of ecologist as public figures in the environmental movement with special knowledge of how natural systems work. Cornell ecologist LaMont Cole is an important figure to examine both of these trends. Like many of his contemporaries, Cole was devoted to a perspective on natural selection operating at levels above the individual. However, because of his influential mathematical treatment of animal demography he has been historically subsumed into a group of scholars that views the events in the life course as adaptations to the maximization of individual fitness--life history theory. Cole's popular writings and lectures, which consumed his later career, extend his scholarly portrayal of natural populations as tending toward stable and homoeostatic equilibrium, with the goal of drawing contrasts with the deviance of rapid human population growth. In both regards, Cole serves as a topical and temporal extension of the well-documented and analyzed ecology of his mentors--Alfred Emerson, Thomas Park, and Warder Allee--in the University of Chicago Zoology Department.     

Effects of predator-induced thinning and activity changes on life history in a damselfly

We investigated how the lethal and non-lethal presence and absence of a fish predator, perch (Perca fluviatils), influenced behaviour, numbers emerging, size at emergence, and development rate of the damselfly Lestes sponsa. The experiment was carried out in outdoor artificial ponds and spanned from the egg stage to emergence of the damselflies. During the experiment food resources for the damselflies were continuously monitored. Damselflies exposed to a lethal predator showed a significantly lower activity level than those in the absence of predators or subjected to a non-lethal predator. Half-way through the larval stage the reduction in activity level was correlated with the presence of lethal predators, and at the end of the larval stage with higher zooplankton densities. Though larvae decreased activity level, size at emergence was larger and development time faster for individuals in the lethal predator treatment. Since fewer larvae emerged from that treatment we interpret the larger size at emergence to be an effect of a combination of thinning and higher zooplankton densities.     

Management implications of fish trap effectiveness in adjacent coral reef and gorgonian habitats

A combination of visual census and trap sampling in St. John, USVI indicated that traps performed better in gorgonian habitat than in adjacent coral reef habitat. Although most families were seen more commonly in coral habitat, they were caught more often in gorgonian areas. Traps probably fished more effectively in gorgonian habitats, especially for migrating species, because traps provided shelter in the relatively topographically uniform environment of gorgonian dominated habitats. Recently, trap fishermen on St. John have been moving effort away from traditionally fished nearshore coral reefs and into a variety of more homogeneous habitats such as gorgonian habitat. Consequently, exploitation rates of the already over-harvested reef fish resources may be increasing. Reef fish managers and marine reserve designers should consider limiting trap fishing in gorgonian habitats to slow the decline of reef fisheries.     

Chemistry,histochemistry and microscopy of the organic matrix of spicules from a gorgonian coral

Summary Alcian blue dye normally binds to polyanionic, polymeric substances. Such structures are often associated with calcium binding portions of the organic matrix in calcifying tissues. The organic matrix of spicules prepared from the gorgonian Pseudoplexaura flagellosa (Houttuyn) is alcianophilic. The dye is very tightly bound to the lipoid portion of the insoluble spicule matrix. No acidic substances (sulfated or acidic polysaccharides or phospholipids) were demonstrable in this material, suggesting an unustial but unknown interaction between dye and substrate. On a microscopical basis, inclusion of Alcian blue (or Ruthenium red) is an essential co-requisite to glutaraldehyde fixation. Without the dye the morphological integrity of the spicule is lost on decalcification. The fragmented matrix is still alcianophilic suggesting that the dye may substitute for material solubilized by the decalcifying agents. Examination of post-decalcification supernatants demonstrate that approximately 13% of the matrix is solubilized on demineralization, releasing 93% of the carbohydrate but <20% of the protein. Liberated protein takes the form of peptides ranging from 1100–1500 daltons. The composition of these peptides is a function of the demineralizing agent. Acidic demineralizers produce peptides proportionately high in acidic amino acids, that do not bind calcium. Peptides produced by chelator decalcification appear to bind calcium but other evidence strongly suggests that the binding is due to adsorbed chelator rather than by soluble matrix.     

Leaf age and life history variables of a leafminer: the case of Liriomyza trifolii on potato leaves

Field observations and laboratory experiments have demonstrated that infestation by the serpentine leaf miner, Liriomyza trifolii Burgess (Diptera: Agromyzidae), begins in the lower leaves of the potato plant, and proceeds through the middle leaves to those of the upper canopy. In choice and no‐choice experiments, mated adult L. trifolii females were given access for 48 h to potato plants, and caged on differently aged leaves. The extent of their feeding and oviposition on the 5–7 leaflets of the upper, middle, and lower leaves were recorded. The life history variables of the next generation were estimated: percentage egg hatch, number of mines formed, larval survival, number of pupae formed, size and weight of pupae, percentage pupation, number of adults formed, percentage emergence, size and weight of adults, sex ratio, adult longevity, and their reproductive performance. The results showed that L. trifolii females laid fewer eggs on the upper leaves, which were poor hosts for larvae. However, a comparison of oviposition behavior between the middle and lower leaves showed that the data did not fit the oviposition preference–offspring performance hypothesis, which postulates that females preferentially oviposit on hosts on which larvae perform best. Females exhibited a preference for the larger, older, lower leaves, although the middle leaves were superior for the growth and development of the young stages. It is hypothesized that adult ovipositional preference for the older, larger, and thicker leaves of the lower foliage may be influenced by factors other than resource quality for larvae.     

Population diversity in salmon: linkages among response,genetic and life history diversity

Response diversity and asynchrony are important for stability and resilience of meta‐populations, however little is known about the mechanisms that might drive such processes. In salmon populations, response diversity and asynchrony have been linked to the stability of their meta‐populations and the fisheries that integrate across them. We examined how population diversity influenced response diversity and asynchrony in 42 populations of Chinook salmon from the Fraser River, British Columbia. We examined diversity in the survival responses to large‐scale ocean climate variables for populations that differed in life history. Different life‐histories responded differently to ocean environmental conditions. For instance, an increase of offshore temperature was associated with decreased survival for a population with ocean rearing juveniles but increased survival for a population with stream rearing juveniles. In a second analysis, we examined asynchrony in abundance between populations, which we then correlated with life history, spatial, and genetic diversity. Populations that were more genetically distant had the most different population dynamics. Collectively, these results suggest that fine‐scale population diversity can contribute to the asynchrony and response diversity that underpins the stability of fisheries or metapopulation dynamics, and emphasize the need to manage and conserve this scale of population diversity.     

Population history and life history influence the migration rate of female Glanville fritillary butterflies

This study examines the causes of emigration from small fragments of suitable habitat in a species that has a distinct metapopulation structure, frequent turnover of local populations, and substantial migration among local populations and currently unoccupied habitat fragments. We conducted a field experiment in which 727 individuals of the Glanville fritillary butterfly ( Melitaea cinxia ) originating from four regions were marked and released simultaneously in a natural environment. In three of the four source regions, larvae for the experiment were collected from dozens of small local populations, some of which had been established in the previous summer (new populations), whereas the remaining populations were older. In two of the source regions, female butterflies prefer a host plant ( Veronica spicata ) that is not present in the release area, where there is only Plantago lanceolata , the preferred host plant of females from the other two source regions. We found that migration rate of males was unrelated to any of the factors studied in this experiment. In contrast, two factors influenced the migration rate of females. First, Veronica -preferring females had higher emigration rate than Plantago- preferring females from the Plantago -containing release patches, demonstrating that the individual perception of habitat quality significantly influences the migration rate of females. Second, females from newly-established populations were more dispersive than females from older populations, supporting the notion that metapopulation processes (recurrent colonizations) select for increased migration. The observed migration rate was not correlated with any body size measurements, and thus the observed differences in migration rate were apparently caused by differences in the behaviour of female butterflies rather than in their flight capacity.     

Predator-induced life history changes in amphibians: egg predation induces hatching

The timing of transitions between life history stages should be affected by factors that influence survival and growth of organisms in adjacent life history stages. In a series of laboratory experiments, we examined the influence of predation risk as a cue to trigger a life history switch in amphibians. In the Oregon Cascade Mountains, some populations of Pacific treefrogs ( Hyla regilla ) and Cascades frogs ( Rana cascadae ) are under intense egg predation by predatory leeches (families Glossiphonidae and Erpobdellidae). We document that both treefrogs and Cascades frogs show plasticity in hatching characteristics in response to the threat of egg predation. Pacific treefrogs hatch sooner and at an earlier developmental stage when either predatory leeches or non-predatory earthworms are allowed direct contact with the developing egg mass. The same response is elicited even without direct contact. Chemical cues of predatory leeches and chemicals released from injured eggs appear to elicit the same early hatching response in treefrogs. For Cascades frogs, cues of leeches, but not those of injured eggs, elicit an early hatching response. Hatching early in response to egg predators may reduce predation. Plasticity of hatching characteristics has rarely been examined. However, we suspect that it may be common, particularly in populations or species that experience high variability in predation pressure between years.     

From inducible defences to population dynamics: modelling refuge use and life history changes in Daphnia

We investigated the relative importance of a behavioural defence (refuge use through diel vertical migration) and a life history change (a reduced size at first reproduction) that are used by daphnids to decrease the risk of predation by visually hunting fish. We used an individual based model of a Daphnia population in a stratified lake to quantify the effects of these inducible defences on Daphnia predation-mortality and the resulting Daphnia population dynamics. Our analysis shows that diel vertical migration (DVM) confers a much stronger protection against fish predation than a reduced size at first reproduction (SFR). DVM allows daphnids to withstand a higher predation pressure in the epilimnion and it decelerates a Daphnia population decline more strongly than a reduced SFR. DVM effectively reduces the (P/B) flow of carbon from daphnids to fish.
Many theoretical studies have only considered the fitness benefits of DVM above 'staying up' in the epilimnion of a lake. Our results suggest that 'staying down' in the hypolimnion would confer an even stronger fitness benefit to Daphnia than DVM at times of peak predation risk. Daphnids that remain in the hypolimnion avoid the predation suffered by migrating daphnids around dusk and dawn. Staying down could prevent a Daphnia population decline, while DVM and a reduced SFR can only decelerate the decrease of Daphnia population densities under heavy fish predation. Staying down at high concentrations of fish infochemicals has in fact been observed within a variety of Daphnia clones and species, both in the laboratory and in stratified lakes.