Food limitation in a wild cyclopoid copepod population: direct and indirect life history responses

We studied the effects of food limitation on the population dynamics of the freshwater cyclopoid copepod Diacyclops thomasi in Oneida Lake, New York. In the field population, maximum juvenile abundance coincided seasonally with high phytoflagellate concentration. During the clear-water phase (a seasonal period of low algal density), D. thomasi disappeared from the water column, but fourth-instar copepodids (CIV) were found encysted in developmental arrest in the sediment. Laboratory assays of the effect of the density of two types of food on copepod life history parameters showed that temporal variation in the concentration of relatively small phytoflagellates significantly affected stage-specific development times. This food limitation was most pronounced during the clear-water phase, and supplementation of the diet with a laboratory-cultured phytoflagellate, Chlamydomonas, prevented food limitation. Although developmental arrest appears to be controlled primarily by photoperiod, availability of the larger, more mobile food, Euglena, also influenced the percentage of individuals entering developmental arrest in the laboratory. An investigation of the spatial and temporal emergence pattern in the field revealed that CIV copepodids started to emerge in late autumn and that emergence rates were significantly greater at deep-water locations (9–12 m water depth) compared with shallow-water locations (5–7 m). The clear-water phase in Oneida Lake is an annual event, probably produced by intense grazing by Daphnia pulicaria and Daphnia galeata. Food limitation is thus very likely a recurrent phenomenon for D. thomasi. This apparent seasonal competitive impact of Daphnia on Diacyclops affects both nauplii and immature copepodids. Diacyclops shows two types of responses to the food limitation: (1) the physiological response of slowed active development, and (2) the adaptive response of developmental arrest. Received: 3 November 1997 / Accepted: 1 March 1998     

Condition and size of damselflies: a field study of food limitation

Summary Based on evidence from field manipulations, several authors have recently suggested that interference competition among larval odonates reduces individual growth rates and biomass by reducing foraging rates. This study was designed to test the effects of food shortage on condition (relative mass per unit head width) of larval Ischnura verticalis (Odonata: Coenagrionidae) under laboratory conditions and to use these results to estimate the degree of food shortage of larvae under naturally occurring field conditions. In the laboratory, there were marked differences in condition of larvae fed diets ranging from ad libitum feeding with worms to ad libitum feeding with Daphnia 1 day out of every 8. Condition of larvae collected from May through October from 17 different sites in southern Ontario indicated that, for most of the year, larvae had conditions similar to those fed ad libitum with Daphnia in the laboratory. There was no evidence that larval condition was related to population density. Condition of larvae in most sites during July was similar to that of larvae fed poor diets in the laboratory. It is unlikely that the low conditions were due to competition as there were no correlations with density across sites and population densities during July were at their lowest. Adult head widths showed a seasonal decline from mid June to the end of the flight season. There was no evidence that head widths were related to population density although there was some evidence that head widths of males were positively related to larval condition. My results do not support the hypothesis that competition is important in affecting foraging rates and subsequent development of larvae. Contrasts between my results and other studies may stem from difficulties with the interpretation of field experiments, that densities in my study may have been low due to fish predation, and/or that I. verticalis larvae are slow moving relative to other larvae and thus less likely to interact.     

Experimental studies of exploitative competition in a grazing stream insect

Summary Field and laboratory experiments were conducted to determine whether intraspecific competition for food occurs during the larval stage of the periphyton-grazing caddisfly Glossosoma nigrior (Trichoptera: Glossosomatidae). Larvae were placed in field enclosures at densities less than, equal to, or greater than their natural densities. Most of these individuals began to pupate after 3 weeks, whereupon the mass of each individual was determined. Final mass declined significantly as larval densities increased, whereas neither developmental rate nor mortality/emigration rate was significantly affected by density manipulations. a supplemental experiment comparing the final mass of individuals grown at reduced densities in a laboratory stream with individuals from a natural stream bottom confirmed the results of the more extensive field experiment: reductions in density resulted in significant increases in final mass. Periphyton availability in field enclosures declined according to a negative exponential function as larval densities increased. Over the 25-fold range of larval densities used in these experiments, the final mass of individuals increased linearly with periphyton standing crops. This result suggests that Glossosoma larvae may compete for food even at densities below those employed in this study. Path analysis was used to explore the importance of indirect (i.e., exploitative) and direct (i.e, interference) mechanisms for the observed competitive effects. The analysis indicates that a model based solely on exploitation explains nearly as much of the variance in mass as a model incorporating both interference and exploitation.     

Optimal conservation effort for a population in a stochastic environment

We study the optimal conservation effort for a population in a fluctuating environment. The survivorship of a population is affected by unpredictable environmental fluctuation (noise) and can be improved by conservation effort accompanied by a cost. The optimal effort level is the one that minimizes the total cost, defined as the weighted sum of the population extinction risk and the economic cost of conservation effort. The optimal effort depends on the variance and the probability distribution of the noise, the relative importance of the population's survival vs. the economic cost, the effectiveness of conservation effort, and the time scope over which we optimize. The analysis of dynamic programming illustrates that the choice of extinction risk function greatly affects the optimal effort level. The conservation effort level that is the best solution of a multiple-year optimization may be higher than that for the corresponding single-year optimization, if the population is relatively safe. However, the conservation level for the multiple-year optimization becomes lower than for the single-year optimization if the population is endangered. In a similar manner, the optimal conservation effort level for the problem with a short time scope is either higher or lower than that for the problem with a long time scope, depending on the extinction risk of the population. Next, for each parameter of the model, we define five different sensitivities of extinction probability or of the total cost. We then study the mean increase in the total cost caused by the uncertainty of parameters. To achieve the best conservation result, we need to invest the limited research effort to the parameter with the largest effect to the optimal effort level, rather than to those with large impacts on the extinction probability or on the total cost. The recommended policy should depend critically on the choice of the criterion to optimize, which shows the importance of theoretical study of the relationship in performing proper decision making in conservation practice.     

Resource limitation and intraspecific patterns of weight x length variation among spring detritivores

This study examined the intraspecific variability of a phenotypic trait, the body weight/body length ratio, and its adequacy to provide unbiased information about patterns of resource availability among conspecific individuals. Individual body weight and length were measured for the amphipod Gammarus minus Say (and other detritivores) in samples from freshwater springs differing in expected resource availability, and from sites in which detritus abundance had been manipulated. Mean individual weight per length was lower: (a) in populations from low-richness than from high-richness detritivore guilds; (b) in populations with size-abundance distributions similar to that of the entire guild, rather than statistically different; (c) in population samples from outside than from inside the areas of detritus addition. Small-sized individuals (< 3 mm) showed the largest variation among both populations and population samples. Similar differences were qualitatively observed for the other common detritivores co-occurring with G. minus in some springs. These observed patterns were in agreement with the variation of resource availability expected among field conditions, supporting the relevance of weight per length as a measure of food limitation.     

Relative importance of competition with Daphnia (Cladocera) and nutrient limitation on Anuraeopsis (Rotifera) population dynamics in a laboratory study

• 1 In the laboratory, the growth and reproduction of Anuraeopsis fissa were measured when fed on Scenedesmus species grown in nutrient‐sufficient, nitrogen‐limited and phosphorus‐limited media and in the presence or absence of one adult Daphnia longispina per vial.
• 2 Poor food quality may reduce the effect of competition on rotifers. Competition from Daphnia was stronger with nutrient sufficient algae than with nutrient limited algae. P‐limitation significantly reduced Anuraeopsis population growth rate and fecundity. The effect of nutrient limitation on Anuraeopsis was stronger than that of competition with Daphnia. The Anuraeopsis population declined with P‐limited food in both the presence and absence of Daphnia.
• 3 Exploitative competition by Daphnia on Anuraeopsis was stronger in the nutrient‐sufficient treatment than in the N‐limited one. Density, fecundity and population growth rate of Anuraeopsis were negatively affected by Daphnia in the nutrient‐sufficient treatment, while only fecundity was reduced by Daphnia in the N‐limited treatment. Consequently, in the N‐limited treatment, mortality should be lower in the presence of Daphnia. This result could suggest that Anuraeopsis lives longer when short of nitrogen.
• 4 Nutrient limitation may affect to the competitive interactions between zooplankton species. P‐limitation decreased the quality of algae as food for Anuraeopsis while N‐limitation decreased the susceptibility of this rotifer species to exploitative competition by Daphnia.

     

Interspecific competition among grasshoppers and their effect on plant abundance in experimental field environments

Summary We tested whether grasshoppers in experimental field environments, i.e. cages (40×40 cm) placed on existing old field vegetation, (1) were limited in density by plant abundance and/or nitrogen content, (2) exhibited interspecific competition, and (3) altered the relative abundance of different plant species. We examined interactions among a pair of early season grasshopper species (May–June; Arphia conspersa and Pardalophora apiculata) and a late season pair (July–August; Melanoplus femur-rubrum and Melanoplus bivittatus). Each grasshopper species was placed in cages by itself and with another grasshopper species. Grasshoppers generally survived at higher density in fertilized cages and they reduced plant abundance relative to empty cages, suggesting that grasshoppers may be food limited at these densities. In unfertilized plots, early season grasshoppers preferred grasses (Schizachyrium scoparium and Poa pratensis) and favored the growth of forbs (especially Solidago spp.). However, late in summer, Melanoplus spp. preferred Solidago spp. and favored the growth of grasses.The pattern of grasshopper survivorship and plant reduction within these experimental environments provide preliminary support for some of the predictions of resource competition theory. Grasshoppers exhibited interspecific competition only if they significantly reduced plant biomass. If two species competed, a grasshopper species was eliminated only if the superior competitor, when living by itself, could reduce plant biomass to a significantly lower level than the inferior competitor. Competitors persisted only if they did not differ in their ability to reduce plant biomass or reduced the abundance of different plant species.     

Phytoplankton growth in the Ohio, Cumberland and Tennessee Rivers, USA: inter-site differences in light and nutrient limitation

Seasonal patterns in resource limitation of phytoplankton growth were assessed monthly within three large rivers with differing extents of water regulation. The Ohio River is regulated by low dams that do not substantially modify discharge, while the Cumberland and Tennessee Rivers are impounded by a series of high dams to enhance water storage for downstream flood control. Laboratory dilution assays with light and nutrient manipulations indicated that light was the main factor limiting phytoplankton growth at irradiances below 7 E m^–2 d^–1. Light limited growth was frequent in the turbid, higher discharge of the Ohio River, but was rare in the heavily regulated Tennessee and Cumberland Rivers. When irradiance exceeded 7 E m^–2 d^–1, phytoplankton were either P-limited (Cumberland River), co-limited by P and N (Tennessee River), or Si limited (Ohio River). Site-specific differences in nutrient limitation were consistent with differences in ambient nutrient levels, with the Tennessee and Cumberland Rivers characterized by lower N and P concentrations, and the Ohio River by lower Si. Downstream nutrient depletion was evident in the Ohio River through comparison of an upstream and a downstream site, with nutrient limitation (Si) occurring more frequently downstream. Phytoplankton growth rates at ambient light and nutrient levels ranged from 0.1 to 1.5 d^–1 in the Ohio River and 0.2 to 0.6 d^–1 in the Tennessee and Cumberland Rivers. Growth rates were greatest at the onset of the summer base pool, as light intensities increased and nutrient levels were maximal. Our findings indicate that multiple factors regulate phytoplankton growth in regulated rivers and that spatial complexity may arise from differences in discharge and water aging.     

Role of nutrient limitation in the competition between uropathogenic strains of Klebsiella pneumoniae and Escherichia coli in mixed biofilms

Klebsiella pneumoniae and Escherichia coli form mixed species biofilms in catheter-associated urinary tract infections. Recently, a detrimental effect of K. pneumoniae over E. coli was observed in mixed species biofilms grown in an artificial urine medium. The mechanism behind this competitive interaction was studied. K. pneumoniae partially outcompeted E. coli in early-stage batch-fed biofilms, whereas both microorganisms co-exist at longer times (K. pneumoniae:E. coli ratio, 55:1), as shown by cell counts and confocal microscopy. E. coli cells were scattered along the K. pneumoniae biofilm. Biofilm supernatants did not appear to contain either antimicrobial or anti-biofilm activities against E. coli. Biofilms grown under continuous flow prevented interspecies competition. K. pneumoniae showed both increased siderophore production and better growth in iron-limited media compared to E. coli. In summary, these results indicate the importance of nutrient (particularly iron) competition in the modulation of the bacterial composition of mixed species biofilms formed by uropathogenic K. pneumoniae and E. coli.     

Predation, competition, and nutrient recycling: a stoichiometric approach with multiple nutrients

A model for two competing prey species and one predator is formulated in which three essential nutrients can limit growth of all populations. Prey take up dissolved nutrients and predators ingest prey, assimilating a portion of ingested nutrients and recycling or respiring the balance. For all species, the nutrient contents of individuals vary and growth is coupled to increasing content of the limiting nutrient. This model was parameterized to describe a flagellate preying on two bacterial species, with carbon (C), nitrogen (N), and phosphorus (P) as nutrients. Parameters were chosen so that the two prey species would stably coexist without predators under some nutrient supply conditions. Using numerical simulations, the long-term outcomes of competition and predation were explored for a gradient of N:P supply ratios, varying C supply, and varying preference of the predator for the two prey. Coexistence and competitive exclusion both occurred under some conditions of nutrient supply and predator preference. As in simpler models of competition and predation these outcomes were largely governed by apparent competition mediated by the predator, and resource competition for nutrients whose effective supply was partly governed by nutrient recycling also mediated by the predator. For relatively small regions of parameter space, more complex outcomes with multiple attractors or three-species limit cycles occurred. The multiple constraints posed by multiple nutrients held the amplitudes of these cycles in check, limiting the influence of complex dynamics on competitive outcomes for the parameter ranges explored.     

Alternation of bottom-up and top-down regulation in a natural population of an agromyzid leafminer,Chromatomyia suikazurae

The population dynamics and the relative importance of bottom-up and top-down effects in a plant-leafminer-multiparasitoid interaction was studied between 1981 and 1990 in a natural forest in Kyoto, Japan. The leafminer, Chromatomyia suikazurae (Agromyzidae, Diptera), passed two generations (G1 and G2) on Lonicera gracilipes (Caprifoliaceae). The G1 population in February was free from parasitoid attack, and the mortality in G1 was mainly caused by resource limitation. Intraspecific competition for resources occurred at the larval stage in G1, and the larval mortality was density-dependent. The G1 adult density was resource-limited (the number of newly opened leaves), and its variability was lower than that of G2. The G2 population in April was not resource-limited but subject to intense attack by a species-rich parasitoid complex, and thus total mortality was much larger than that in G1. Significant density dependence was detected not in larval but in pupal mortalities, which were mainly caused by parasitism by parasitoids that attacked the pupa. The host population alternately experienced bottom-up effects during the larval stage in G1 and top-down effects during the pupal stage in G2. Overall population fluctuation was non-cyclic and mainly due to climatically-induced fluctuation of available plant resources in G1.     

Simultaneous effects of food limitation and inducible resistance on herbivore population dynamics

Many herbivore populations fluctuate temporally, but the causes of those fluctuations remain unclear. Plant inducible resistance can theoretically cause herbivore population fluctuations, because herbivory may induce plant changes that reduce the survival or reproduction of later-feeding herbivores. Herbivory can also simply reduce the quantity of food available for later feeders and this, too, can cause population fluctuations. Inducible resistance and food limitation often occur simultaneously, yet whether they jointly facilitate or suppress herbivore fluctuations remains largely unexplored. We present models that suggest that food limitation and inducible resistance may have synergistic effects on herbivore population dynamics. The population-level response of the food plant to herbivory and the details of how inducible resistance affects herbivore performance both influence the resulting herbivore dynamics. Our results identify some biological properties of plant-herbivore systems that might determine whether or not cycles occur, and suggest that future empirical and theoretical population dynamics studies should account for the effects of both food limitation and inducible resistance.     

The role of male contest competition over mates in speciation

Research on the role of sexual selection in the speciation process largely focuses on the diversifying role of mate choice.In particular,much attention has been drawn to the fact that population divergence in mate choice and in the male traits subject to choice directly can lead to assortative mating.However,male contest competition over mates also constitutes an important mechanism of sexual selection.We review recent empirical studies and argue that sexual selection through male contest competition can affect speciation in ways other than mate choice.For example,biases in aggression towards similar competitors can lead to disruptive and negative frequency-dependent selection on the traits used in contest competition in a similar way as competition for other types of limited resources.Moreover,male contest abilities often trade-off against other abilities such as parasite resistance,protection against predators and general stress tolerance.Populations experiencing different ecological conditions should therefore quickly diverge non-randomly in a number of traits including male contest abilities.In resource based breeding systems,a feedback loop between competitive ability and habitat use may lead to further population divergence.We discuss how population divergence in traits used in male contest competition can lead to the build up of reproductive isolation through a number of different pathways.Our main conclusion is that the role of male contest competition in speciation remains largely scientifically unexplored.     

Population and prehistory I: Food-dependent population growth in constant environments

We present a demographic model that describes the feedbacks between food supply, human mortality and fertility rates, and labor availability in expanding populations, where arable land area is not limiting. This model provides a quantitative framework to describe how environment, technology, and culture interact to influence the fates of preindustrial agricultural populations. We present equilibrium conditions and derive approximations for the equilibrium population growth rate, food availability, and other food-dependent measures of population well-being. We examine how the approximations respond to environmental changes and to human choices, and find that the impact of environmental quality depends upon whether it manifests through agricultural yield or maximum (food-independent) survival rates. Human choices can complement or offset environmental effects: greater labor investments increase both population growth and well-being, and therefore can counteract lower agricultural yield, while fertility control decreases the growth rate but can increase or decrease well-being. Finally we establish equilibrium stability criteria, and argue that the potential for loss of local stability at low population growth rates could have important consequences for populations that suffer significant environmental or demographic shocks.     

Life history consequences of sterol availability in the aquatic keystone species <Emphasis Type="Italic">Daphnia</Emphasis>

The absence of essential biochemical nutrients, such as polyunsaturated fatty acids or sterols, has been considered as a mechanism determining trophic interactions between the herbivore Daphnia and its phytoplankton food source. Here, we experimentally quantify the sensitivity of two Daphnia species to decreasing amounts of dietary sterols by measuring variations in life history traits. The two species Daphnia magna and D. galeata were fed different mixtures of the sterol-containing green alga Scenedesmus obliquus and the sterol-free cyanobacterium Synechococcus elongatus; a higher proportion of Synechococcus in the food is equivalent to a decrease in dietary sterols. To address the significance of sterol limitation, the Daphnia species were also fed Synechococcus supplemented with cholesterol. In both species, somatic and population growth rates, maternal dry mass, the number of viable offspring, and the probability of survival were significantly reduced with the lower availability of sterols. A high correlation between the sterol content of the mixed diet and the somatic and population growth rates was found, and growth on cholesterol-supplemented Synechococcus fitted well into this correlation. Somatic growth of first-clutch neonates grown on 100% Synechococcus exhibited a pattern similar to that of somatic growth of their mothers grown on the different food regimes, which demonstrated the significance of maternal effects for sterol-limited population growth. Daphnia galeata had a twofold higher incipient limiting sterol level than D. magna, which indicated interspecific differences in sterol requirements between the two Daphnia species. The results suggest a strong impact of dietary sterols on life history traits and therefore, population dynamics of the keystone species Daphnia.     

Food limitation of population density in the orb-wed spider,Nephila clavata

Field studies were conducted to clarify whether variation in food availability among habitats influences population density, and whether population density has a negative effect on foraging success in the orb-web spider, Nephila clavata. Lifetime food consumption per individual (i.e., foraging success) strongly correlated with mean body size of adult females and mean fecundity in populations. Also, there was a positive correlation between foraging success and population density. Since foraging success reflected potential prey availability in the habitat, food resource appeared to be a limiting factor for populations in this spider. Mean fecundity per individual correlated with population density of the following year, suggesting that decreased reproduction is a major component of food limitation on population density. Consistent defferences in mean body size between particular sites were observed over years, while such difference was less obvious in density. Thus, ranking of food abundance among habitats seems to be predictable between years. A field experiment revealed that an artificial increase in population density had no negative effect on the feeding rate of individuals, suggesting that intraspecific competition for food is not important in this species.     

Food availability and nest predation influence life history traits in Audouin's gull, Larus audouinii

The effects of food availability and nest predation on several life history traits such as adult survival, dispersal, and reproductive performance were assessed in an Audouin's gull (Larus audouinii) colony during the period 1992–1997. The amounts of fish discarded from trawlers were used as a measure of food availability, and a trawling moratorium which partially overlapped with the breeding season of the gulls was taken into account. The effects of nest predation were assessed in 1994, when a terrestrial predator entered the colony and remained for the whole breeding season preying on both eggs and chicks. Using the moratorium and the predatory event as natural experiments, several hypotheses were tested: (a) food supply would affect breeding performance but not adult survival (independently of age and sex), since gulls are long-lived and adult survival is the most sensitive demographic parameter in their population dynamics; (b) the predator would trigger breeding dispersal (although gulls are mostly philopatric, they are known to abandon their natal colony after breeding failure instigated by events such as this). If breeding dispersal occurs, the rate is expected to be higher in females than in males, and higher in new breeders than in more experienced breeding birds, as is usually recorded in colonial seabirds. Probabilities of resighting and survival were estimated separately, using capture-recapture models. As expected, changes in food availability did not affect adult survival, whereas they influenced egg volume, clutch size, and breeding success. Local adult survival was estimated to be 0.908 (SD = 0.007) for males and females, and it did not change significantly with the age of individuals (range 3–8 years). The predator significantly decreased breeding success, and caused the dispersal of a number of adults probably to breed in another colony; this rate was estimated at an average of 0.10 (SD = 0.02). As expected, inexperienced breeders dispersed significantly more (14%) than more experienced breeders (8%) after the predator event, but dispersal was not sex biased. Recapture probabilities after the predator event suggest that birds that left the colony still had not returned. Results confirm that population dynamics of ground-nesting seabirds are sensitive to terrestrial predation, even when predation caused only a partial breeding failure. Received: 16 July 1998 / Accepted: 16 November 1998     

An assessment of overwinter food limitation in a snowshoe hare population at a cyclic low

Snowshoe hares (Lepus americanus) undergo 8- to 11-year population cycles caused by direct and/or interactive effects of overwinter food shortage and predation. However, the demographic significance of food shortage during cyclic population lows remains unclear. I evaluated the importance of overwinter food limitation to the demography (numbers, age and sex ratios) of low-density hare populations during two winters in Manitoba. Also, I examined whether the hypothesized differences in demography of fed and unfed hare populations could be explained by altered movement patterns or social dynamics. Bimonthly live-trapping revealed that food failed to have a direct long-term effect on the number, or change in number, of hares estimated to be on the three supplemented areas, relative to three control areas. Modest numerical responses to supplementation tended to be short-term (i.e., restricted to winter) and related to pre-supplementation densities, with the study area characterized by the highest hare density displaying the strongest and most consistent response to added food. During winter the percentage of females was remarkably variable among study areas and time periods, but added food may have augmented slightly the proportion of females captured in traps. There tended to be slightly more juveniles on supplemented areas during winter periods, and this effect was strongest during the first winter (1991–1992). I found that immigration rates and percentage of hares that were considered to be transient animals were similar on supplemented and control areas, and that spatial distribution of radio-collared animals on versus off of study areas also was similar. Because the overall effect of food on hare populations was small and short-lived, and could be explained largely by small increases in survival and reproduction, I conclude that the study population was not subject to overwinter food limitation. Received: 22 February 1998 / Accepted: 12 February 1999