Evolution and persistence of obligate mutualists and exploiters: competition for partners and evolutionary immunization

Mutualisms are ubiquitous in nature, as is their exploitation by both conspecific and heterospecific cheaters. Yet, evolutionary theory predicts that cheating should be favoured by natural selection. Here, we show theoretically that asymmetrical competition for partners generally determines the evolutionary fate of obligate mutualisms facing exploitation by third-species invaders. When asymmetry in partner competition is relatively weak, mutualists may either exclude exploiters or coexist with them, in which case their co-evolutionary response to exploitation is usually benign. When asymmetry is strong, the mutualists evolve towards evolutionary attractors where they become extremely vulnerable to exploiter invasion. However, exploiter invasion at an early stage of the mutualism's history can deflect mutualists' co-evolutionary trajectories towards slightly different attractors that confer long-term stability against further exploitation. Thus, coexistence of mutualists and exploiters may often involve an historical effect whereby exploiters are co-opted early in mutualism history and provide lasting 'evolutionary immunization' against further invasion.     

Nutrient-induced competition between two species of marine diatoms

Competition experiments betweenPhaeodactylum tricornutum andSkeletonema costatum showed that even at temperatures higher than 10°C (i.c. 14°C), the development ofSkeletonema can be favoured by adjusting nutrient levels and nutrient ratios. Low NSi ratios were found to favourSkeletonema. Additionally, high NP ratios further enhanced the ability ofSkeletonema to dominate the cuftures. Contrary to some statements in literature, it seems that high concentrations of silicates are more important for the dominance ofSkeletonema costatum in large-scale cultures than just low temperatures. This finding is important with regard to stimulating the blooming ofSkeletonema costatum in natural phytoplankton populations as food for bivalve molluscs.     

Interspecific interference competition between two field cricket species

Species interactions, such as interspecific competition, determine population dynamics and community structure. In a park of Amami Oshima Island, Japan, two field cricket species Teleogryllus occipitalis (Audinet‐Serville) (Gryllidae) and Loxoblemmus equestris Saussure (Gryllidae) are dominant species. We examined the interference competition at the nymph and adult stages in the two crickets under controlled laboratory conditions. Survival and emergence rates of T. occipitalis were not affected by the density of L. equestris, whereas survival and emergence rates of L. equestris decreased as the density of T. occipitalis increased. These results showed that T. occipitalis was competitively superior over L. equestris in species interactions. We discuss inconsistency between the results and coexistence of these two species under field conditions.     

Tradeoffs between functional strategies for resource-use and drought-survival in Mediterranean rangeland species

In environments where light is not a limiting resource such as rangelands and grasslands, there is much disagreement regarding the benefits provided by rapid light capture during the growing season and the species’ ability to withstand drought during the dry period. In this study, we selected four perennial herbaceous species with contrasting resource-use strategies (acquisitive versus conservative), which were transplanted as monocultures into PVC pots to evaluate their species-specific responses to drought. The two main strategies of drought-survival (avoidance versus tolerance) were driven by distinct underlying mechanisms that allow the plant to delay or tolerate water deficit in leaves. On the one hand, plants that produced reduced leaves with lower surface area:mass ratio (lower SLA) exhibited higher values of leaf water potential (LWP) and leaf relative water content (LRWC), which could be associated to a higher ability to delay tissue dehydration in enlarged leaves. Regarding the below-ground compartment, dehydration avoidance was promoted by prolonged elongation rates of thinner roots that allow the plant to increase water uptake and accessibility during the dry period. On the other hand, dehydration tolerance was positively related with progressive foliage senescence under water deficit, which probably favored a longer survival of meristematic basal tissues. The results presented in this study suggest the existence of a trade-off between the traits favoring rapid light-acquisition and those enhancing the ability to delay leaf dehydration. Thus, the species related most closely with a resource-acquisition strategy (Bromus erectus and Potentilla neumanniana) could be considered less efficient to delay leaf dehydration than the others (Carex humilis and Festuca christiani-bernardii), as indicated by their lower values of leaf water potential (LWP) and leaf relative water content (LRWC) under identical conditions of water deficit. Our findings support evidence that there is not a single strategy to effectively cope with drought and reveal the diversity of adaptive mechanisms among coexisting species.     

Association between competition and obligate mutualism in a chemostat

In this paper, we consider a simple chemostat model involving two obligate mutualistic species feeding on a limiting substrate. Systems of differential equations are proposed as models of this association. A detailed qualitative analysis is carried out. We show the existence of a domain of coexistence, which is a set of initial conditions in which both species survive. We demonstrate, under certain supplementary assumptions, the uniqueness of the stable equilibrium point which corresponds to the coexistence of the two species.     

Schooling properties of an obligate and a facultative fish species

Changes in attraction and repulsion indicators, depending on the species and the group size, were explored under controlled conditions. Two species, displaying different schooling behaviours in the wild were observed: the bigeye scad Selar crumenophthalmus and the barred flagtail Kuhlia mugil . In the bigeye scad, the polarity and speed were high and stable, and the nearest neighbour distance ( D _NN) decreased when the group size increased. In contrast, for the barred flagtail, polarity and speed decreased according to the group size, inducing a loss of cohesion and leading to a disorganized school. The D _NN mean was stable whatever the group size and relatively high. This experiment indicated that the ability to polarize is first a species-specific trait, rather than a property emerging from the group and led by the circumstances.     

Association between competition and obligate mutualism in a chemostat

In this paper, we consider a simple chemostat model involving two obligate mutualistic species feeding on a limiting substrate. Systems of differential equations are proposed as models of this association. A detailed qualitative analysis is carried out. We show the existence of a domain of coexistence, which is a set of initial conditions in which both species survive. We demonstrate, under certain supplementary assumptions, the uniqueness of the stable equilibrium point which corresponds to the coexistence of the two species.     

Interspecific competition between two species of the bean weevil

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Contributions from the Entomological Laboratory, Kyoto University, No. 110.     

Specificity of the mutualistic association between actinomycete bacteria and two sympatric species of Acromyrmex leaf-cutting ants

Acromyrmex leaf-cutting ants maintain two highly specialized, vertically transmitted mutualistic ectosymbionts: basidiomycete fungi that are cultivated for food in underground gardens and actinomycete Pseudonocardia bacteria that are reared on the cuticle to produce antibiotics that suppress the growth of Escovopsis parasites of the fungus garden. Mutualism stability has been hypothesized to benefit from genetic uniformity of symbionts, as multiple coexisting strains are expected to compete and, thus, reduce the benefit of the symbiosis. However, the Pseudonocardia symbionts are likely to be involved in Red-Queen-like antagonistic co-evolution with Escovopsis so that multiple strains per host might be favoured by selection provided the cost of competition between bacterial strains is low. We examined the genetic uniformity of the Pseudonocardia symbionts of two sympatric species of Acromyrmex ants by comparing partial sequences of the nuclear Elongation Factor-Tu gene. We find no genetic variation in Pseudonocardia symbionts among nest mate workers, neither in Acromyrmex octospinosus, where colonies are founded by a single queen, nor in Acromyrmex echinatior, where mixing of bacterial lineages might happen when unrelated queens cofound a colony. We further show that the two ant species maintain the same pool of Pseudonocardia symbionts, indicating that horizontal transmission occasionally occurs, and that this pool consists of two distinct clades of closely related Pseudonocardia strains. Our finding that individual colonies cultivate a single actinomycete strain is in agreement with predictions from evolutionary theory on host-symbiont conflict over symbiont mixing, but indicates that there may be constraints on the effectiveness of the bacterial symbionts on an evolutionary timescale.     

No competition for resources between two tick species at their parapatric boundary

Summary Aponomma hydrosauri and Amblyomma limbatum are tick species with the same major host species, the sleepy lizard Trachydosaurus rugosus, but parapatric distributions in South Australia. Near Mt. Mary one species is abruptly replaced by the other over one kilometre. There is substantial overlap in feeding sites on the hosts. This suggested that interspecific competition, during feeding, may be an important mechanism preventing further overlap of the distributions of the two species. However, in the overlap region there was no resource shortage, and each species infested hosts independently of the other species. Also in laboratory experiments at the highest densities encountered in the field, larvae and nymphs of each species attached and engorged equally well whether alone or mixed with the other species. These data refuted the, hypothesis of interspecific competition between the two species while feeding.     

Interspecific competition between two species of Trichogrammatidae III The analytical study on the competition

The experiment was carried out in order to analyze the results of interspecific competition which were mentioned in the previous reports (Sasaba 1964, 1965). The following definite tendencies were obtained and an account was given with regard to the process of competition. (1)T. minutum had a higher reproductive rate than T. japonicum at a lower density of the parent wasps. (2) The parasitic efficiency of T. minutum was high as compared with T. japonicum. (3)T. japonicum was consistently superior to T. minutum in the interspecific competition at larval stage.     

Shell fighting and competition between two hermit crab species in Panama

Summary This article is a study of shell fighting between two intertidal hermit crab species in Panama. Laboratory results showed some cases of high exchange frequencies between Calcinus obscurus and Clibanarius albidigitus when the former occupied poor-quality shells. Exchange frequencies varied considerably between collecting sites, and were always low when the defending Clibanarius came from Venado Beach. Shell exchange frequencies estimated from field experiments were similar to those obtained in the laboratory. Observations on relative shell sizes occupied by both species in areas of sympatry and allopatry failed to provide clear evidence that Calcinus reduced the shell size of Clibanarius or that Clibanarius increased the shell size occupied by Calcinus. Results obtained here differ from those obtained in previous studies (Abrams 1980; Bertness 1981a, b), and these differences are discussed. Although shell fighting may be an important component of the interaction of these species, it is likely that roughly 90% of the competition experienced by each species is intraspecific.     

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.     

Experimental evidence of asymmetrical competition between two species of parasitic copepods

Lepeophtheirus thompsoni and Lepeophtheirus europaensis are two parasitic copepods naturally isolated on their sympatric hosts, i.e. turbot (Psetta maxima L.) and brill (Scophthalmus rhombus L.), respectively They are able to meet, mate and hybridize on turbot experimentally but they are naturally prevented from doing so by a strong host preference when given a choice. Theory suggests that such a pattern is possible, but only under conditions of competition for the resource. In the present study the attachment rates of the two copepods were studied experimentally under various conditions of competition, infectious dose and number of available hosts. The results suggest a greater sensitivity to competition for the generalist species L. europaensis than for the specialist L. thompsoni, which is in agreement with theoretical predictions.     

The cost of sex and competition between cyclical and obligate parthenogenetic rotifers

The ubiquity of sexual reproduction is an evolutionary puzzle because asexuality should have major reproductive advantages. Theoretically, transitions to asexuality should confer substantial benefits in population growth and lead to rapid displacement of all sexual ancestors. So far, there have been few rigorous tests of one of the most basic assumptions of the paradox of sex: that asexuals are competitively superior to sexuals immediately after their origin. Here I examine the fitness consequences of very recent transitions to obligate parthenogenesis in the cyclical parthenogenetic rotifer Brachionus calyciflorus. This experimental system differs from previous animal models, since obligate parthenogens were derived from the same maternal genotype as cyclical parthenogens. Obligate parthenogens had similar fitness compared with cyclical parthenogens in terms of the intrinsic rate of increase (calculated from life tables). However, population growth of cyclical parthenogens was predicted to be much lower: sexual female offspring do not contribute to immediate population growth in Brachionus, since they produce either males or diapausing eggs. Hence, if cyclical parthenogens constantly produce a high proportion of sexual offspring, there is a cost of sex, and obligate parthenogens can invade. This prediction was confirmed in laboratory competition experiments.     

Effect of the removal of cleaner fish on the abundance and species composition of reef fish

The ecological significance of cleaner fish on coral reefs was investigated. I removed all cleaner fish, Labroides dimidiatus, from eight small reefs, measured the subsequent effect on the abundance and species composition of all reef fish after 3 and 6 months, and compared it with eight control reefs with cleaner fish. The removal of cleaner fish had no detectable effect on the total abundance of fish on reefs and the total number of fish species at both times. Multivariate analysis by non-metric multidimensional scaling and ANOSIM pairwise tests based on 191 fish species revealed no effect of cleaners on the community structure of fish. Similar results were obtained using principal components analysis on subsets of the data using the 33 most common fish species and the 15 most abundant species (≥5 individuals per reef ) with both log10 (x + 1) transformed data and with fish numbers standardized for abundance. This study demonstrates that the removal of cleaner fish for 6 months did not result in fish suffering increased mortality nor in fish leaving reefs to seek cleaning elsewhere. Received: 28 October 1996 / Accepted: 7 February 1997     

Enhanced colonization success and competition associated with conspecifics in cleaner fish Labroides dimidiatus juveniles

The cleaner fish Labroides dimidiatus affects the abundance of many fishes, including their juveniles, yet how they affect the post-settlement processes of conspecifics remains poorly known. Using a long-term experiment, where L. dimidiatus were regularly removed from seven patch reefs (removals) for 10 years or left undisturbed (controls) on nine, the effect of conspecifics’ presence on recently settled L. dimidiatus juveniles and the relationship between juvenile and adult abundance on control reefs were examined. Repeated sampling 4–40 months after manipulating L. dimidiatus revealed that the abundance of juveniles on removals, compared with controls reefs, was 72 % (95 % CI = 65–79 %) lower; this effect did not vary over time. Thus, decreased colonization was associated with resident conspecific absence and this effect was present after 4 months. Since other studies found 4 months of cleaner absence was too short to affect client abundance—a factor also known to enhance L. dimidiatus colonization—it was likely not involved here. Over 10 years, individual control reefs were almost always occupied by at least one adult (89–100 % of times sampled); but in summer, when L. dimidiatus colonization was highest, juvenile abundance was negatively related with adult abundance at one of two sites, with the expected number of juveniles decreasing by 67 % (95 % CI = 51–83 %) with each additional adult. This suggests a deleterious effect of adults on juveniles at one site, such as competition. Enhanced colonization of juveniles associated with adult presence may partly explain the relative permanence of fish cleaning stations.     

Interspecific competition between introduced house finch populations and two associated passerine species

Summary House Finches (Carpodacus mexicanus), natives of western North America, have expanded their range in the eastern United States since their 1940 release in New York City. Range and the relation of House Finch population growth to the population dynamics of House Sparrows (Passer domesticus) and Purple Finches (Carpodacus purpureus) were examined, using data from the Breeding Bird Survey and the Christmas Bird Count. The House Finch population grew exponentially throughout its eastern range. Significant negative relationships in population density, relative to spatial and temporal control populations, were found between House Finches and House Sparrows in summer and winter, and between House Finches and Purple Finches in summer. Purple Finch and House Sparrow populations outside of the House Finch range appeared to have no effect on each other throughout the study. Neither changes in 74 weather variables, nor changes in forest, field or developed habitat explained the observed trends in population density. The results indicate that House Finches compete with these two species, but winter migration complicates the picture.