An anatomy of interactions among species in a seasonal world

Mathematical models merging biological and predictable seasonal dynamics were used to simulate four types of organism interactions: competition, prey-predation, mutualism and facilitation. By analysing trajectories for biomass in phase portraits (i.e. species 1 biomass plotted against species 2 biomass) graphically and numerically, we found that each of the four interaction types showed characteristic patterns (fingerprints) in phase space. All the four interaction types could be distinguished, even though their time trajectories were strongly modified by seasonal forces. For each of the interaction types, we were able to identify characteristics of the interaction that most strongly distinguished it from the others. We could also assess the relative effect of species characteristics and seasonality on each of the four interactions. The simulations indicate that prey-predation is strongly influenced by seasonal forces and by the characteristics of the predator and its prey. A system variability index got a high value (SVI)=0.167 relative to those of the other interaction types (competition 0.01, mutualism 0.007 and facilitation 0.005). The result was obtained by calculating, angles between successive vectors linking pairs of samples and the positive x-axis and arranging these as frequency histograms for each of the four quadrants of the phase portrait. In an effort to capture circular motions and other characteristics of the trajectories sampled, angle frequencies were analysed using multivariate statistics (PCA), yielding characteristic directions in phase space. We believe that the characteristic patterns identified also will be found in real time series.     

Experimental small-scale grassland fragmentation alters competitive interactions among ant species

Different species may respond differently to habitat fragmentation. Theory predicts that abundant generalist species should be less affected by fragmentation than specialist species. In ant communities, the most abundant species is often behaviourally dominant. Thus, habitat fragmentation could alter competitive interactions between the dominant ant species and the other species. We tested this hypothesis in a long-term grassland fragmentation experiment. Fragments of different size (20.25 and 2.25 m²) were isolated by a 5-m wide strip of frequently mown vegetation. Control plots were situated in adjacent undisturbed grassland. Ant density and species composition were assessed 3 and 6 years after initiation of the experimental fragmentation. The effect of the dominant ant species on the resource use of the other species was examined at natural sugar resources (aphids and extrafloral nectaries) and at artificial sugar baits. Lasius paralienus was the most abundant ant species (72% of nests) in the grasslands examined. Species richness and forager density in the other species decreased with increasing density of L. paralienus in fragments but not in control plots. The overall forager density of the other species was positively related to their habitat niche overlap with L. paralienus. The density of foragers of the other species at sugar resources was not affected by L. paralienus forager density. The experimental fragmentation resulted in an increase in natural sugar resources in fragments. This may have reduced the intensity of interspecific competition for sugar resources. Our study shows that the grassland fragmentation altered interactions between the dominant L. paralienus and the other ant species.Electronic Supplementary Material Supplementary material is available for this article at     

On selection dynamics for competitive interactions

In this paper, we are interested in an integro-differential model that describe the evolution of a population structured with respect to a continuous trait. Under some assumption, we are able to find an entropy for the system, and show that some steady solutions are globally stable. The stability conditions we find are coherent with those of Adaptive Dynamics.     

Variation and seasonal changes in hoverfly species: interactions between temperature, age and genotype

Eristalis tenax L. and E. intricarius L. are two hoverflies which vary considerably in colour pattern. Whilst much of the phenotypic variation in both species is due to genetic variation at major gene loci, there are interactions with pupal temperature and with age of adult. Low pupal temperatures produce, on average, darker abdominal patterns in E. tenax , although the effect is hard to pick out in natural populations. Changes of pattern with age convert a bimodal autumn population to a unimodal post-hibernation population in the following spring. Hair colour is also made darker by cold treatment. Pupal treatments have a strong effect on hair colouration in E. intricarius. High temperatures inhibit the production of black hairs on the thorax, but not all the genotypes are equally sensitive to temperature. Seasonal fluctuations in colour pattern frequencies are detectable in E. intricarius. In three Eristalis species that have been studied so far, the interactions between genotype, age and temperature operate in quite different ways. The temperature responses may be relevant to theories of thermal melanism, although the colour pattern polymorphisms are, more obviously, examples of Batesian mimicry.     

Endogenous timing in competitive interactions among relatives

Most evolutionary game theory models solve for equilibrium levels of some behaviour on the restrictive assumptions that players choose their actions simultaneously, and that a player cannot change its action after observing that of its opponent. An alternative framework is provided by sequential or 'Stackelberg' games in which one player commits to a 'first move' and the other has an opportunity to observe this move before choosing its response. Recent interest in the economic literature has focused on Stackelberg games which exhibit 'endogenous timing', i.e. games in which a leader and a follower arise spontaneously as a consequence of each player attempting to maximize its reward. Here, we provide the first demonstration of endogenous timing in an evolutionary context using a simple model of resource competition (the 'tug-of-war' model). We show that whenever two related individuals compete for a share of communal resources, both do best to adopt distinct roles in a sequential game rather than engage in simultaneous competition. Somewhat counterintuitively, the stable solution is for the weaker individual to act as leader and commit to a first move, because this arrangement leads to a lower total effort invested in competition. Endogenous timing offers a new explanation for the spontaneous emergence of leaders and followers in social groups, and highlights the benefits of commitment in social interaction.     

An analysis of seasonal dynamics of a mixed population of Daphnia, and the associated cladoceran community

SUMMARY. The dynamics of a mixed population of Daphnia (D. ambigua and D. parvula) are analysed over 9 months in a small eutrophic lake. A spring peak of D. ambigua occurred shortly after ice break-up. Decline of D. ambigua coincided with a high occurrence of sexual reproduction, the appearance of D. parvula which did not become abundant, and a substantial increase in Ceriodaphnia quadrangula . A third cladoceran, Bosmina longirostris , was abundant before and after the period of numerical dominance by D. ambigua , but declined to very low numbers coincident with the increase in D. ambigua.
Estimates of egg hatching rate at various temperatures demonstrate that these three cladocerans have different rate responses to temperature, and that seasonal succession of species is predictable from temperature alone. The importance of temperature versus species interactions in controlling species succession is discussed.     

Resource partitioning among three woodpecker species Dendrocopos spp. during the breeding season

Food composition, prey size utilization and foraging behaviour of three sympatric woodpecker species ( Dendrocopos major, D. medius, D. minor ) were studied in an oak forest near Budapest during the breeding season in 1983 and 1984. Considering these three aspects of feeding, the great spotted woodpecker is a generalist species. Food composition of this species resembled the arthropod supply on the bark of trees more than those of the other two species. The bark of the trees seems to be a relatively unproductive microhabitat in the breeding season, so woodpecker species use, to different degrees, the food supply of the foliage as well. The food and the foraging behaviour of the middle spotted woodpecker show that this species feeds on prey living both on barks and in the foliage; it occupies-an intermediate position between the great and the lesser spotted woodpeckers. Prey size did not correlate with predator size suggesting that woodpeckers adapted not to the summer resources but rather the winter ones.     

Ice-cover effects on competitive interactions between two fish species

1. Variations in the strength of ecological interactions between seasons have received little attention, despite an increased focus on climate alterations on ecosystems. Particularly, the winter situation is often neglected when studying competitive interactions. In northern temperate freshwaters, winter implies low temperatures and reduced food availability, but also strong reduction in ambient light because of ice and snow cover. Here, we study how brown trout [Salmo trutta (L.)] respond to variations in ice-cover duration and competition with Arctic charr [Salvelinus alpinus (L.)], by linking laboratory-derived physiological performance and field data on variation in abundance among and within natural brown trout populations. 2. Both Arctic charr and brown trout reduced resting metabolic rate under simulated ice-cover (darkness) in the laboratory, compared to no ice (6-h daylight). However, in contrast to brown trout, Arctic charr was able to obtain positive growth rate in darkness and had higher food intake in tank experiments than brown trout. Arctic charr also performed better (lower energy loss) under simulated ice-cover in a semi-natural environment with natural food supply. 3. When comparing brown trout biomass across 190 Norwegian lakes along a climate gradient, longer ice-covered duration decreased the biomass only in lakes where brown trout lived together with Arctic charr. We were not able to detect any effect of ice-cover on brown trout biomass in lakes where brown trout was the only fish species. 4. Similarly, a 25-year time series from a lake with both brown trout and Arctic charr showed that brown trout population growth rate depended on the interaction between ice breakup date and Arctic charr abundance. High charr abundance was correlated with low trout population growth rate only in combination with long winters. 5. In conclusion, the two species differed in performance under ice, and the observed outcome of competition in natural populations was strongly dependent on duration of the ice-covered period. Our study shows that changes in ice phenology may alter species interactions in Northern aquatic systems. Increased knowledge of how adaptations to winter conditions differ among coexisting species is therefore vital for our understanding of ecological impacts of climate change.     

On detecting interactions between species in population dynamics

If we are to progress out of our current state of uncertainty about the role of interspecific competition in community structure, it is essential that we can distinguish competition from other types of population interaction, and from lack of interaction, in particular case studies. To make such distinctions, it is necessary to quantify the effect of species on each other. One way to do this is to calculate interaction coefficients, and another (only readily applicable in experimental systems) is to plot graphs of N against time and to contrast monocultures with mixed cultures. We show (1) that these methods often appear to give contradictory results, and (2) that the problems are most pronounced when one species has a low equilibrium population size in mixed culture. To resolve the question of whether an interspecific interaction is taking place (and if so, what kind of interaction), it is necessary to apply tests of significance which overcome the problems of serial correlation inherent in all long-term population experiments. We illustrate the use of such tests in the analysis of the results from an experimental Drosophiia system. In the past, this kind of test has generally not been applied, and this raises the question of whether some of the 'classic cases of competition' in the experimental literature were really competition at all.     

Analysis of three species models of mutualism in predator-prey and competitive systems

Mutualism arises in many qualitatively different ways, but previous 2-species models of mutualism correspond to only a few of these. Following Addicott and Freedman (1983), we present two models of mutualism in which interactions among three species lead to mutualism between two of them. One model involves interactions among a predator, a mutualist-prey, and a mutualist, while the other involves interactions among a competitor, a mutualist-competitor, and a mutualist. Given biologically reasonable constraints upon the functions in the model, we present (1) the conditions for boundedness of solutions, (2) the equilibria and their local stability, and (3) the conditions for the existence of small amplitude periodic solutions, a behavior not predicted by 2-species models of mutualism. The models include costs to the mutualist-prey or mutualist-competitor of associating with the mutualist. Analysis of special cases shows that mutualism can cause the extinction of the predator, or the reversal of competitive outcomes.     

An analysis of competitive interactions between 3 hermit crab species

Summary Competition for empty gastropod shells in a group of three sympatric hermit crabs (Pagurus hirsutiusculus, Pagurus granosimanus, and Pagurus beringanus) was studied in the San Juan Archipelago, Washington State. Estimates of the competitive effects of each species on the others' shell supplies were derived using field data on shell utilization and the results of laboratory experiments to determine rates of acquisition and exchange of shells and preferences for different shell species. Each species experienced approximately an order of magnitude more intraspecific competition than interspecific competition for empty shells. This resulted from differences in preference for shell shapes, shell size use, and habitat use between P. hirsutiusculus and P. granosimanus, and largely from differences in habitat use between P. beringanus and the other two species. Experiments involving the release and recensusing of marked empty shells were used to estimate competitive effects more directly for the interaction between P. hirsutiusculus and P. granosimanus. Results were consistent with the estimates derived from data on resource partitioning. Possible causes of the low levels of interspecific competition are discussed, and results are compared with studies of other organisms that estimated both inter- and intra-specific competition.     

Qualitative dynamics of three species predator-prey systems

Summary A qualitative analysis of some two and three species predator-prey models is achieved by application of the method of averaging in conjunction with a Lyapunov function constructed from the appropriate Volterra-Lotka model. We calculate the limit cycle solution for a two-species model with a Holling type functional response of the predator to its prey by means of a time-scaled transformation. The existence of a bifurcation of steady states for a community of three species is discussed and the periodic solution around one of the unstable steady states is calculated to the lowest approximation. Several comments are made regarding the behavior of these systems under changes of some control parameters.This work was supported in parts by USERDA, Contract number E(11-1)-3001.     

Ontogenetic shifts in competitive interactions and intra-guild predation between two wolf spider species

Abstract. 1. The wolf spiders (Araneae: Lycosidae) Hogna helluo (Walckenaer) and Pardosa milvina Hentz co-occur in soybean fields of south-west Ohio, U.S.A. As adults, Hogna is the larger species and has the competitive advantage in most interactions; due to differing phenologies, however, their size-classes frequently overlap and as such there is potential for shifts in competitive ability and intra-guild predation. The hypothesis that competitive interactions and intra-guild predation will favour Pardosa when Pardosa is similar-sized, or has a size advantage over Hogna , was tested in laboratory and field experiments.
2. Studies in laboratory arenas, pairing similar-sized individuals of these species and Hogna spiderlings with larger spiders of both species, revealed that intra-guild predation seldom occurs with similar-sized Hogna and Pardosa , however Pardosa will consume small Hogna individuals in laboratory arenas.
3. Field experiments involved stocking high densities (50 m^{– 2} ) of Pardosa and Hogna in enclosures placed in soybean fields. In experiments with spiders of similar size, no interspecific effects were uncovered, but an intraspecific effect was found for Pardosa as its survival and weight gain were lower in the presence of more conspecifics. Large Hogna or Pardosa had no effect on the survival or weight gain of Hogna spiderlings.
4. Although Hogna is a better competitor as an adult, it has no advantage over Pardosa when their size-classes overlap, and Pardosa effects on Hogna may be inconsequential under field conditions. Therefore, the co-existence of these species is fostered by the fact that there are few negative interspecific interactions during their ontogeny.     

The dynamics of photosynthetic acclimation to changes in light quanlity and quality in three Australian rainforest tree species

Photosynthetic acclimation was studied in seedlings of three subtropical rainforest species representing early (Omalanthus populifolius), middle (Duboisia myoporoides) and late (Acmena ingens) successional stages in forest development. Changes in the photosynthetic characteristics of pre-existing leaves were observed following the transfer of plants between deep shade (1–5% of photosynthetically active radiation (PAR), selectively filtered to produce a red/far-red (R/FR) ratio of 0.1) and open glasshouse (60% PAR and a R/FR ratio of 1.1–1.2), and vice versa. The extent and rate of response of the photosynthetic characteristics of each species to changes in light environment were recorded in this simulation of gap formation and canopy closure/overtopping. The light regimes to which plants were exposed produced significant levels of acclimation in all the photosynthetic parameters examined. Following transfer from high to low light, the light-saturated rate of photosynthesis was maintained near pre-transfer levels for 7 days, after which it decreased to levels which closely approximated those in leaves which had developed in low light. The decrease in photosynthetic capacity was associated with lower apparent quantum yields and stomatal conductances. Dark respiration was the parameter most sensitive to changes in light environment, and responded significantly during the first 4–7 days after transfer. Acclimation of photosynthetic capacity to increases in irradiance was significant in two of the three species studied, but was clearly limited in comparison with that of new leaves produced in the high light conditions. This limitation was most pronounced in the early-successional-stage species, O. populifolius. It is likely that structural characteristics of the leaves, imposed at the time of leaf expansion, are largely responsible for the limitations in photosynthetic acclimation to increases in irradiance.     

Effect of the epizoic rotifer Brachionus rubens on the population growth of three cladoceran species

Using population densities and growth rates as criteria, we studied interactions between the epizoic rotifer Brachionus rubens and each of three cladoceran species differing in size and reproductive rates — Daphnia carinata, Moina macrocopa and Ceriodaphnia rigaudi. In all mixed — species experiments, B. rubens existed in both the epizoic mode, attached to the cladoceran host, and in the free-swimming mode. Rotifer population growth rates were significantly depressed in the presence of M. macrocopa, presumably as a consequence of exploitative and interference competition. The largest cladoceran, D. carinata probably did not suppress B. rubens, because the epizoic component of the rotifer population escaped from the deleterious effects of mechanical interference. Peak population numbers and initial population growth rates reached by all three cladocerans were lower in the presence of B. rubens, probably because of the adverse effects of the epizoic infestation, which was maximal on D. carinata and least on C. rigaudi. In mixed-species cultures of D. carinata and M. macrocopa, the presence of B. rubens helped D. carinata coexist with M. macrocopa, which otherwise would have suppressed the Daphnia.     

Complex food webs prevent competitive exclusion among producer species

Herbivorous top-down forces and bottom-up competition for nutrients determine the coexistence and relative biomass patterns of producer species. Combining models of predator-prey and producer-nutrient interactions with a structural model of complex food webs, I investigated these two aspects in a dynamic food-web model. While competitive exclusion leads to persistence of only one producer species in 99.7% of the simulated simple producer communities without consumers, embedding the same producer communities in complex food webs generally yields producer coexistence. In simple producer communities, the producers with the most efficient nutrient-intake rates increase in biomass until they competitively exclude inferior producers. In food webs, herbivory predominantly reduces the biomass density of those producers that dominated in producer communities, which yields a more even biomass distribution. In contrast to prior analyses of simple modules, this facilitation of producer coexistence by herbivory does not require a trade-off between the nutrient-intake efficiency and the resistance to herbivory. The local network structure of food webs (top-down effects of the number of herbivores and the herbivores' maximum consumption rates) and the nutrient supply (bottom-up effect) interactively determine the relative biomass densities of the producer species. A strong negative feedback loop emerges in food webs: factors that increase producer biomasses also increase herbivory, which reduces producer biomasses. This negative feedback loop regulates the coexistence and biomass patterns of the producers by balancing biomass increases of producers and biomass fluxes to herbivores, which prevents competitive exclusion.     

Factors regulating cladoceran dynamics in a Venezuelan floodplain lake

Five cladoceran zooplankton species in a Venezuelan floodplainlake, Laguna la Orsinera, were analyzed to identify factorscontrolling their dynamics over a period encompassing isolation,inundation by the Orinoco River and drainage. The abundanceof each species increased rapidly to a maximum associated withflooding of the lake by the Orinoco River. We calculated birth,death and population growth rates, and developed a method forestimating uncertainty in these parameters, in order to interpretobserved cladoceran abundance patterns. Rapid increases in populationsizes resulted both from high birth rates associated with decreasingturbidity following inundation and from the hatching of restingeggs. Birth rates remained high throughout the study, but highdeath rates restricted each population to a brief maximum andthen maintained small population sizes throughout the remainderof the study. Our demographic analyses suggest that high mortalityresulted from intense predation by fish and by Chaoborus larvae,rather than from resource limitation, and that this mortalitywas the major factor regulating cladoceran abundance duringinundation and drainage in this tropical floodplain lake.     

Phytoplankton of Lake Peipsi-Pihkva: species composition, biomass and seasonal dynamics

With 33 years of phytoplankton quantitative studies carried out, a series of qualitative data with a length of over 80 years is at our disposal. About 500 algal species have been found in plankton by different researchers. In different seasons and years 35 main species (dominants and subdominants) form 68–96 % of biomass in L. Pihkva (southern, more eutrophic part) and 60–97 % in L. Peipsi (northern, less eutrophic part). L. Lämmijärv, connecting the two parts is similar to L. Pihkva in respect to phytoplankton and the trophic state. Diatoms and blue-green algae prevail in biomass, diatoms and green algae, in the species number. The oligo-mesotrophic Aulacoseira islandica (O. Müller) Sim. is characteristic of the cool period; A. granulata (Ehr.) Sim. and Stephanodiscus binderanus (Kütz.) Krieger prevail in summer and autumn, the latter being most abundant in the southern part. Gloeotrichia echinulata (J.S. Smith) P Richter and Aphanizomenon flos-aquae (L.) Ralfs dominate in summer causing water-bloom. Phytoplankton has mostly three maxima in seasonal dynamics in L. Peipsi and two in L. Pihkva. Its average biomass in spring in different years has fluctuated in the range 5.6–16 and 6–12.7 g m^–3, in summer 3.1–14.8 and 5.6–125 (10–20 in most cases); and in autumn 7–16.3 and 5.2–26 in the northern and southern parts, respectively.The dominant complex has not changed considerably since 1909; however, the distribution of dominant species in lake parts has become more even in the last decades. Periods of high biomass occurred in the first half of the 1960s and 1970s and in 1988–1994, of low biomass in 1981–1987. The first coincided, in general, with periods of low water level and high water temperature.     

Variation in mycorrhizal growth response influences competitive interactions and mechanisms of plant species coexistence

Plant species vary in their growth response to arbuscular mycorrhizal (AM) fungi, with responses ranging from negative to positive. Differences in response to AM fungi may affect competition between plant species, influencing their ability to coexist. We hypothesized that positively responding species, whose growth is stimulated by AM fungi, will experience stronger intraspecific competition and weaker interspecific competition in soil containing AM fungi, while neutrally or negatively responding species should experience weaker intraspecific and stronger interspecific competition. We grew Plantago lanceolata, which responds positively to AM fungi, and Bromus inermis, which responds negatively to AM fungi, in an additive response surface competition experiment that varied the total density and relative frequency of each species. Plants were grown in sterilized background soil that had been inoculated with whole soil biota, which includes AM fungi, or a microbial wash, that contained other soil microbes but no AM fungi, or in sterilized soil that contained no biota. The positively responding P. lanceolata was more strongly limited by intraspecific than interspecific competition when AM fungi were present. By contrast, the presence of AM fungi decreased the strength of intraspecific competition experienced by the negatively responding B. inermis. Because AM fungi are almost always present in soil, strong intraspecific competition in positively responding species would prevent them from outcompeting species that respond neutrally or negatively to AM fungi. The potential for increased intraspecific competition to offset growth benefits of AM fungi could, therefore, be a stabilizing mechanism that promotes coexistence among plant species.