Are Trade-Offs Among Species’ Ecological Interactions Scale Dependent? A Test Using Pitcher-Plant Inquiline Species

Trade-offs among species' ecological interactions is a pervasive explanation for species coexistence. The traits associated with trade-offs are typically measured to mechanistically explain species coexistence at a single spatial scale. However, species potentially interact at multiple scales and this may be reflected in the traits among coexisting species. I quantified species' ecological traits associated with the trade-offs expected at both local (competitive ability and predator tolerance) and regional (competitive ability and colonization rate) community scales. The most common species (four protozoa and a rotifer) from the middle trophic level of a pitcher plant (Sarracenia purpurea) inquiline community were used to link species traits to previously observed patterns of species diversity and abundance. Traits associated with trade-offs (competitive ability, predator tolerance, and colonization rate) and other ecological traits (size, growth rate, and carrying capacity) were measured for each of the focal species. Traits were correlated with one another with a negative relationship indicative of a trade-off. Protozoan and rotifer species exhibited a negative relationship between competitive ability and predator tolerance, indicative of coexistence at the local community scale. There was no relationship between competitive ability and colonization rate. Size, growth rate, and carrying capacity were correlated with each other and the trade-off traits: Size was related to both competitive ability and predator tolerance, but growth rate and carrying capacity were correlated with predator tolerance. When partial correlations were conducted controlling for size, growth rate and carrying capacity, the trade-offs largely disappeared. These results imply that body size is the trait that provides the basis for ecological interactions and trade-offs. Altogether, this study showed that the examination of species' traits in the context of coexistence at different scales can contribute to our understanding of the mechanisms underlying community structure.     

Egg size, offspring quality and food level in planktonic rotifers

1. For two species of planktonic rotifers, Brachionus calyciflorus and Synchaeta pectinata , neonates from larger eggs were more resistant to starvation than neonates from smaller eggs, so egg size was positively correlated with offspring quality.
2. Theories of optimal offspring size in zooplankton predict either a decrease in offspring size with increasing food level, or maximum offspring size at intermediate food levels. In experiments using acclimated, even-aged individuals, neither rotifer species changed its egg size as predicted by theory. B . calyciflorus egg size increased with increasing food concentration, while S . pectinata egg size did not change.
3. The lack of fit to theory may have occurred because of constraints on minimum or maximum egg size, constraints on total reproductive effort, or because offspring fitness is influenced by traits in addition to neonate starvation time.     

Evolution of Rotifer Life Histories

When compared to most other multicellular animals, rotifers are all relatively small, short-lived and fast-reproducing organisms. However among and within different rotifer species there is a large variation in life history patterns. This review accounts for such variation in rotifers, with a strong focus on monogonont rotifers. As the life cycle of monogonont rotifers involves both asexual and sexual reproduction, life history patterns can be examined on the level of the genetic individual, which includes all asexual females, sexual females and males that originated from one resting egg. This concept has been applied successfully in many areas, for example in predicting optimal levels of mictic reproduction or sex allocation theory. The benefits and implications of the view of the genetic individual are discussed in detail. Rotifer life histories can also be viewed on the level of physiological individuals. A large part of this review deals with the life histories of individual amictic females and addresses life history traits like body size, egg size and resource allocation patterns. It asks which trade-offs exist among those traits, how these traits change under the influence of environmental factors like food availability or temperature, and whether these changes can be interpreted as adaptive.     

A minimalist approach to the effects of density-dependent competition on insect life-history traits

Abstract  1. Due to its effects on the phenotypic and genotypic expression of life-history traits, density-dependent competition is an important factor regulating the growth of populations. Specifically for insects, density-dependent competition among juveniles is often associated with increased juvenile mortality, delayed maturity, and reduced adult size.
2. The aim of the work reported here was to test whether the established phenotypic effects of density-dependent competition on life-history traits could be reproduced in an experimental design requiring a minimal number of individuals. Larvae of the mosquito Aedes aegypti were reared at densities of one, two, or three individuals per standard Drosophila vial and in six different conditions of larval food availability. This design required relatively few individuals per independent replicate and included a control treatment where individuals reared at a density of one larva per vial experienced no density-dependent interactions with other larvae.
3. Increased larval densities or reduced food availability led to increased larval mortality, delayed pupation, and the emergence of smaller adults that starved to death in a shorter time (indicating emergence with fewer nutritional reserves).
4. Female mosquitoes were relatively larger than males (as measured by wing length) but males tended to survive for longer. These differences increased as larval food availability increased, indicating the relative importance of these two traits for the fitness of each sex. The role of nutritional reserves for the reproductive success of males was highlighted in particular.
5. This minimalist approach may provide a useful model for investigating the effects of density-dependent competition on insect life-history traits.     

Phenotypic plasticity and constancy of life-history traits in laboratory clones of Daphnia magna Straus: effects of neonatal length

1. The phenotypic constancy of four laboratory Daphnia magna clones in fitness-related life-history traits, such as age and clutch size at maturity, was studied among consecutive experimental runs in differing food environments.
2. A significant part of the observed clonal and genetic-by-environmental variation in age and clutch size at maturity was explained by experimentally uncontrollable variations in neonatal body length.
3. Despite food availability, neonatal length determined the number of instars invested to maturity and thus maturation age. Clonal differences in neonatal length and thus in maturation instar occurrence across environments explained most of the clonal variability observed in maturation age.
4. Although interclonal differences in clutch size existed, most of the phenotypic plasticity observed for clutch size was mediated by clonal differences in neonatal length.
5. Clonal differences in neonatal length and in the occurrence of maturation instars across environments dramatically affected the body length of instar IM-2 where provisioning of eggs take place. Since clutch size is determined from clutch mass and clutch mass was strongly related to the body length of instar IM-2, clonal differences across environments in body length of instar IM-2 mirrored clonal differences across environments in clutch size.
6. The results reported in the present study show that maternally mediated traits such as neonatal length affect how genotypes respond to different environmental selection regimes (genetic-by-environmental interaction). Future research needs to focus on the effects of neonatal length on the heritability or genetic variation of the reaction norms, since prediction of the response to selection is a key research objective in quantitative genetic studies.     

Population growth rates of two closely related rotifer species: effects of food quantity, particle size, and nutritional quality

SUMMARY. 1. The relationship between population growth rates and the concentrations of several algal species was determined in laboratory experiments with the rotifers Brachionus rubens and B. calyciflorus .
2. The effects of food quantity were well described by a modified Monod model with a threshold for zero population growth. The model parameters depended on particle size and nutritional quality of the food algae. Differences between the rotifer species were significant and reflected their varying food-size preferences.
3. For each rotifer species, thresholds were lowest for algae in the most readily ingested size range. The lowest thresholds were 0.07–0.09 mgC 1⁻¹ with algae of about 5 μm equivalent spherical diameter (ESD) for B. rubens , and 0.19 mgC 1⁻¹ with algae of about 10 μm ESD for B. calyciflorus .
4. Maximal growth rates ( r _max) were slightly below 0.8 day⁻¹ for both rotifers with most algal species. The highest r _max values for both rotifers were observed when Cyclotella meneghiniana was provided as food. With this alga, B. calyciflorus had a significantly higher r_max (1.02 day⁻¹) than B. rubens (0.838 day⁻¹).
5. From a comparison of the relationship between growth rates and ingestion rates, Chlamydomonas reinhardii appeared to be of low nutritional quality for B. rubens .
6. Egg ratios were related to growth rate and were not influenced by the algal food used. Egg development times and average mortality rates were estimated from the relationship between egg ratio and growth rate. B. calyciflorus appeared to have a high average mortality rate (0.383 day⁻¹) compared to B. rubens (0.083 day⁻¹).     

Ecophysiological traits may explain the abundance of climbing plant species across the light gradient in a temperate rainforest

Climbing plants are a key component of rainforests, but mechanistic approaches to their distribution and abundance are scarce. In a southern temperate rainforest, we addressed whether the dominance of climbing plants across light environments is associated with the expression of ecophysiological traits. In mature forest and canopy gaps, we measured leaf size, specific leaf area, photosynthetic rate, and dark respiration in six of the most abundant woody vines. Mean values of traits and their phenotypic change (%) between mature forest and canopy gaps were predictor variables. Leaf size and specific leaf area were not significantly associated with climbing plant dominance. Variation in gas-exchange traits between mature forest and canopy gaps explained, at least partly, the dominance of climbers in this forest. A greater increase in photosynthetic rate and a lower increase in dark respiration rate when canopy openings occur were related to the success of climbing plant species. Dominant climbers showed a strategy of maximizing exploitation of resource availability but minimizing metabolic costs. Results may reflect phenotypic plasticity or genetic differentiation in ecophysiological traits between light environments. It is suggested that the dominant climbers in this temperate rainforest would be able to cope with forest clearings due to human activities.     

Life-history variation in closely related generalist predators living in the same habitat: a case study with three Cyclosa spiders

1. Generalist predators sharing similar food resources and phenologies as well as having no competitive interactions are expected to have a similar life-history pattern, but some closely related web spiders show different life-history traits. The present paper clarifies possible selection pressures affecting life-history traits of the three coexisting Cyclosa spiders and explores the significance of the life-history variation.
2. Cyclosa argenteoalba had lower daily survival rate and higher growth rate, C. sedeculata had higher daily survival rate and lower growth rate, and C. octotuberculata showed intermediate levels. This implies that the selection pressures these spiders experience differ appreciably even in the same habitat.
3. The significance of the life-history characteristics of the three species was evaluated by general life-history theories. Cyclosa argenteoalba showed distinguishing reproductive traits: shorter time to maturation, larger reproductive effort, larger relative clutch size, decreased clutch size with the number of clutches, and smaller egg size. These characteristics may have evolved in response to the larger ratio of juvenile to adult survivorship. Cyclosa octotuberculata had a clutch size much larger than the other two species, but relative clutch sizes accounting for body size were similar between C. octotuberculata and C. sedeculata . Also, the two species showed a similar time to maturation despite having different selection pressures. Probably, higher growth rate compensates for lower survivorship, leading to the similarity in some reproductive traits.     

Fertile Crescent crop progenitors gained a competitive advantage from large seedlings

1. Cereal domestication during the transition to agriculture resulted in widespread food production, but why only certain species were domesticated remains unknown. We tested whether seedlings of crop progenitors share functional traits that could give them a competitive advantage within anthropogenic environments, including higher germination, greater seedling survival, faster growth rates, and greater competitive ability.
2. Fifteen wild grass species from the Fertile Crescent were grown individually under controlled conditions to evaluate differences in growth between cereal crop progenitors and other wild species that were never domesticated. Differences in germination, seedling survival, and competitive ability were measured by growing a subset of these species as monocultures and mixtures.
3. Crop progenitors had greater germination success, germinated more quickly and had greater aboveground biomass when grown in competition with other species. There was no evidence of a difference in seedling survival, but seed size was positively correlated with a number of traits, including net assimilation rates, greater germination success, and faster germination under competition. In mixtures, the positive effect of seed mass on germination success and speed of germination was even more beneficial for crop progenitors than for other wild species, suggesting greater fitness. Thus, selection for larger seeded individuals under competition may have been stronger in the crop progenitors.
4. The strong competitive ability of Fertile Crescent cereal crop progenitors, linked to their larger seedling size, represents an important ecological difference between these species and other wild grasses in the region. It is consistent with the hypothesis that competition within plant communities surrounding human settlements, or under early cultivation, benefited progenitor species, favoring their success as crops.

     

Clonal variation in the survival and reproduction of Daphniapulicaria under low-food stress

1. Survivorship and reproductive parameters of eight Daphnia pulicaria clones were evaluated in a life-table experiment under four dietary regimes ranging from food limitation to complete food deprivation.
2. D . pulicaria exhibited both diet-independent interclonal differences and genotype–environment interactions in life-span.
3. Reproduction ceased entirely or nearly so under the three treatments with lowest food availability. Under life-long food limitation, differences between clones were evident in clutch sizes and age at first reproduction. Some clones produced ephippia at a markedly higher frequency than others under food-limiting conditions.
4. For most of the life-history traits evaluated, within-lake variation among clones exceeded differences between the two lake populations they represented.
5. Intraspecific variations in response to periods of food deprivation and to extended quantitative or qualitative food-limitation could affect the clonal composition of Daphnia populations. They are therefore of potential importance in determining the effect of altered phytoplankton assemblages on zooplankton communities.     

Clonal variation in the survival and reproduction of Daphniapulicaria under low-food stress

1. Survivorship and reproductive parameters of eight Daphnia pulicaria clones were evaluated in a life-table experiment under four dietary regimes ranging from food limitation to complete food deprivation.
2. D . pulicaria exhibited both diet-independent interclonal differences and genotype–environment interactions in life-span.
3. Reproduction ceased entirely or nearly so under the three treatments with lowest food availability. Under life-long food limitation, differences between clones were evident in clutch sizes and age at first reproduction. Some clones produced ephippia at a markedly higher frequency than others under food-limiting conditions.
4. For most of the life-history traits evaluated, within-lake variation among clones exceeded differences between the two lake populations they represented.
5. Intraspecific variations in response to periods of food deprivation and to extended quantitative or qualitative food-limitation could affect the clonal composition of Daphnia populations. They are therefore of potential importance in determining the effect of altered phytoplankton assemblages on zooplankton communities.     

High-dimensional coexistence of temperate tree species: functional traits, demographic rates, life-history stages, and their physical context

Theoretical models indicate that trade-offs between growth and survival strategies of tree species can lead to coexistence across life history stages (ontogeny) and physical conditions experienced by individuals. There exist predicted physiological mechanisms regulating these trade-offs, such as an investment in leaf characters that may increase survival in stressful environments at the expense of investment in bole or root growth. Confirming these mechanisms, however, requires that potential environmental, ontogenetic, and trait influences are analyzed together. Here, we infer growth and mortality of tree species given size, site, and light characteristics from forest inventory data from Wisconsin to test hypotheses about growth-survival trade-offs given species functional trait values under different ontogenetic and environmental states. A series of regression analyses including traits and rates their interactions with environmental and ontogenetic stages supported the relationships between traits and vital rates expected from the expectations from tree physiology. A combined model including interactions between all variables indicated that relationships between demographic rates and functional traits supports growth-survival trade-offs and their differences across species in high-dimensional niche space. The combined model explained 65% of the variation in tree growth and supports a concept of community coexistence similar to Hutchinson's n-dimensional hypervolume and not a low-dimensional niche model or neutral model.     

Direct and indirect impact of two common rotifer species (Keratella spp.) on two abundant ciliate species (Urotricha furcata, Balanion planctonicum)

1. We investigated the potential competition and feeding impact of the common rotifer species, Keratella cochlearis and K. quadrata , on the abundant prostomatid ciliates, Urotricha furcata and Balanion planctonicum, in laboratory batch culture experiments. All four species have similar feeding preferences, co-occur in many freshwater environments, and are thus potential competitors for the same algal food.
2. Two small Cryptomonas species served as food for the ciliates and the rotifers in the experiments. Growth rates of each ciliate species were measured when they grew alone and when they were paired with one of the rotifer species.
3. Both rotifer species reduced the growth rate of U. furcata , probably primarily by direct feeding on the ciliates. Growth rate of B. planctonicum was unaffected by K. cochlearis , but was drastically reduced by grazing and/or mechanical interference of K. quadrata .
4. These results suggest niche partitioning of the sympatric ciliates with respect to their rotifer competitors/predators.     

Root plasticity of native and invasive Great Basin species in response to soil nitrogen heterogeneity

Soil nutrients are heterogeneously distributed in natural systems. While many species respond to this heterogeneity through root system plasticity, little is known about how the magnitude of these responses may vary between native and invasive species. We quantified root morphological and physiological plasticity of co-occurring native and invasive Great Basin species in response to soil nitrogen heterogeneity and determined if trade-offs exist between these foraging responses and species relative growth rate or root system biomass. The nine study species included three perennial bunchgrasses, three perennial forbs, and three invasive perennial forbs. The plants were grown in large pots outdoors. Once a week for 4 weeks equal amounts of ¹⁵NH₄ ¹⁵NO₃ were distributed in the soil either evenly through the soil profile, in four patches, or in two patches. All species acquired more N in patches compared to when N was applied evenly through the soil profile. None of the species increased root length density in enriched patches compared to control patches but all species increased root N uptake rate in enriched patches. There was a positive relationship between N uptake rate, relative growth rate, and root system biomass. Path analysis indicated that these positive interrelationships among traits could provide one explanation of how invasive forbs were able to capture 2 and 15-fold more N from enriched patches compared to the native grasses and forbs, respectively. Results from this pot study suggest that plant traits related to nutrient capture in heterogeneous soil environments may be positively correlated which could potentially promote size-asymmetric competition belowground and facilitate the spread of invasive species. However, field experiments with plants in different neighbor environments ultimately are needed to determine if these positive relationships among traits influence competitive ability and invader success.     

Polyploidy and invasion success: trait trade-offs in native and introduced cytotypes of two Asteraceae species

Invasion success is favoured by the introduction of pre-adapted genotypes. In addition, novel pressures in the introduced range may lead to phenotypic changes related to fitness or competitive ability of introduced plants. Polyploidy appears to be over-represented in invasive plants, but differences between cytotypes in growth strategies including trade-offs among plant traits have received little attention so far in the context of biological invasions. We grew Centaurea stoebe L. and Senecio inaequidens D.C. in a greenhouse experiment to test for differences in fitness (shoot biomass, reproductive output) and competitive ability (vegetative size, specific leaf area, leaf dry matter content, root–shoot ratio) between diploid and polyploid cytotypes as well as between native and introduced plants. For both species, diploid and tetraploid genotypes occur in the native range, whereas only tetraploids are present in the introduced range. In the native range of both species, diploid and tetraploid genotypes had different growth strategies. Tetraploid genotypes of C. stoebe and S. inaequidens had, respectively, higher specific leaf area and stem height than diploid ones. Thus, for both species, native tetraploids appeared more competitive than native diploids, which could explain, at least partially, the invasion success of the pre-adapted tetraploid genotypes. The comparison of native and introduced tetraploid genotypes revealed differences in traits linked to competitive ability, which could be linked to novel selection in the new environment. In S. inaequidens, we found evidence for a competition-colonisation trade-off, whereas persistence of C. stoebe in the new range seemed to be linked to a competition-defence trade-off.     

Feeding strategies in drosophilid parasitoids: the impact of natural food resources on energy reserves in females

1. Adult feeding strongly increases longevity and fecundity in parasitic wasps. Searching for food resources involves costs; therefore, it is advantageous to be able to feed on the breeding substrate.
2. Cohorts of females of four drosophilid parasitoid species were assigned to different food treatments including starvation. Fat contents were measured and compared after 5 days. In this way the nutritional value of breeding substrates to females could be expressed in terms of energy reserves.
3. Three of the four species tested fed on natural breeding substrates. A fourth species did not produce evidence of such feeding. However, this species fed on honey when fat reserves were low.
4. Feeding is related to: (i) the species' natural breeding substrate, (ii) the energy allocation and lifetime expectancy of the female, and (iii) time of the season.
5. Because feeding strongly influences longevity and fecundity, it is argued that the different feeding strategies found may affect competitive relationships between these drosophilid parasitoid species.     

Asynchronous temporal variation among sites in condition of two carabid species

Abstract.  1. It is generally assumed that generalist predatory carabids are food limited, and the degree of food limitation can vary in space and time, but there are only a few studies that investigate whether or not temporal variation in insect condition is synchronised among sites.
2. In this paper temporal and spatial variation in four fitness-related traits – body mass corrected for size, body size, fat content, and fecundity – of two carabid species, Pterostichus cupreus (L.) and Pterostichus melanarius (Ill.), differing in overwintering strategy, lifespan, diet, and habitat breadth were studied.
3. Interactions between temporal and spatial variation for the fitness-related traits were found for both species. Temporal changes in food limitation were not synchronised among sites, and fitness-related traits were not generally higher on some sites than on others.
4. There was an effect of study year on body mass, fat content, and fecundity for both species, suggesting that temporal environmental variation over large areas is more important than the spatial environmental variation within a year.
5. Within a time period, most different fitness-related traits, except body size, were positively correlated to each other among individuals of the same species. The fitness-related traits of females and males of the same species were generally positively correlated. However, despite the fact that the food resources of the two species overlap, there was no correlation in fitness-related traits between the two species.     

Intraspecific trait variation and trade‐offs within and across populations of a toxic dinoflagellate

Intraspecific trait diversity can promote the success of a species, as complementarity of functional traits within populations may enhance its competitive success and facilitates resilience to changing environmental conditions. Here, we experimentally determined the variation and relationships between traits in 15 strains of the toxic dinoflagellate Alexandrium ostenfeldii derived from two populations. Measured traits included growth rate, cell size, elemental composition, nitrogen uptake kinetics, toxin production and allelochemical potency. Our results demonstrate substantial variation in all analysed traits both within and across populations, particularly in nitrogen affinity, which was even comparable to interspecific variation across phytoplankton species. We found distinct trade‐offs between maximum nitrogen uptake rate and affinity, and between defensive and competitive traits. Furthermore, we identified differences in trait variation between the genetically similar populations. The observed high trait variation may facilitate development and resilience of harmful algal blooms under dynamic environmental conditions.     

Consumption of cyanobacteria by roach (Rutilus rutilus): useful or harmful to the fish?

1. The ability of roach to use cyanobacterial food is generally believed to be one reason for the dominance of roach over perch in eutrophic European lakes. The aim of this study was to test whether cyanobacteria really are a suitable food for juvenile roach. Special attention was paid to differences between the two cyanobacteria species Aphanizomenon and Microcystis which are common in eutrophic lakes and are ingested by roach there.
2. We performed growth and behaviour experiments with juvenile roach fed with zooplankton and the different cyanobacteria. Growth rate with Aphanizomenon was lower than with Daphnia but significantly higher than without food, whereas growth rate with Microcystis was as low as without food.
3. In cultivation experiments of roach faeces, Microcystis was found not to have been digested and grew exponentially after passing through the gut whereas Aphanizomenon stayed at low biomass. Differences in growth were not related to the toxin content of cyanobacteria. Investigations of roach motility showed no differences whether fed with Aphanizomenon or Microcystis .
4. In contrast to Microcystis , Aphanizomenon can be regarded as a suitable food source for juvenile roach probably because of its better digestability. We conclude that the ability to feed on cyanobacteria is not a general competitive advantage for roach, but the outcome depends on the species composition of the cyanobacteria.     

Increasing returns in the life history of Columbian ground squirrels

1. We examined positive associations and trade-offs of maternal and reproductive traits in a population of Columbian ground squirrels, Spermophilus columbianus .
2. Structural size, body condition, mother's personal allocation to body mass during reproduction, and timing of littering were estimated for live-trapped reproductive females that were observed during an 8-year period, and were compared to litter mass, litter size, and average pup mass using path analyses.
3. Mothers exhibited age-structured traits that influenced reproductive patterns. Yearling mothers were significantly smaller, bred later, and had smaller litters than older females. Mothers that gained more body mass during reproduction and older mothers in good body condition that were structurally large had larger litters.
4. Early seasonal timing of littering was an important positive influence on successful reproduction by older mothers only in early breeding seasons and in years when conditions for reproduction were good for all females.
5. The number of offspring that survived to 1 year of age was most strongly associated with litter mass and litter size; date of breeding was of secondary influence, with earlier litters exhibiting greater success.
6. In general, mothers that gained the most in body mass during reproduction were concurrently more successful in weaning large litters (perhaps due to better quality of foraging habitat).
7. In addition to expected reproductive trade-offs, reproduction by Columbian ground squirrels exhibited positive associations of life-history traits that may reflect evolutionary increasing returns.