Experimental analysis of the evolutionary potential of hybridization in leopard frogs (Anura: Ranidae)

Artificial crossing using Rana blairi and R. sphenocephala frogs produced conspecific, interspecific and F₁ backcross hybrid genotypes. Although hybrid males used in the crosses were sterile, crosses using hybrid females produced viable larvae. The larval performance of resultant parental and hybrid genotypes was measured in experimental ponds at two densities. Density significantly affected survival, body mass at metamorphosis, larval period length and metamorphosis for all genotypes. Survival was the same among genotypes, but decreased with increasing density. Body mass at metamorphosis was the same among genotypes, but decreased with increasing density. Larval period increased with increasing density. Among genotypes, larvae from the conspecific R. sphenocephala cross had the shortest larval period while larvae from the conspecific R. blairi cross had the longest larval period. All hybrid genotypes had larval periods longer than R. sphenocephala, but shorter than R. blairi. The percentage of individuals metamorphosing was highest for R. sphenocephala ponds and lowest for R. blairi ponds across densities. Ponds with hybrid larvae produced a greater proportion of metamorphs than those with R. blairi larvae, but a smaller proportion than R. sphenocephala ponds. Equivalent or increased relative larval performance of hybrid genotypes under the conditions of our experiment suggests that hybrid genotypes may possess similar or higher fitnesses relative to their progenitors in some environments. Reduced fertility of adult hybrid males is a powerful selective force against natural hybridization. Nevertheless, because of the successful reproduction by female hybrids, natural hybridization has the potential to serve as a mechanism for the introgression of novel genetic variation that can benefit both R. blairi and R. sphenocephala in fluctuating and unpredictable larval environments. Experimental determination of the fitness of parental and hybrid genotypes is crucial for a comprehensive understanding of the effects of hybridization on organismal evolution.     

HYBRIDIZATION IN LEOPARD FROGS (RANA PIPIENS COMPLEX): LARVAL FITNESS COMPONENTS IN SINGLE-GENOTYPE POPULATIONS AND MIXTURES

Recognizing the predominant mode of selection in hybrid systems is important in predicting the evolutionary fate of recombinant genotypes. Natural selection is endogenous if hybrid genotypes are at a disadvantage relative to parental species independent of environment. Alternatively, relative fitness can vary in response to environmental variation (exogenous selection), and hybrid genotypes can possess fitness values equal to or greater than that of parental species. I investigated the nature of natural selection in a leopard frog hybrid system by rearing larvae of hybrid and parental genotypes between Rana blairi and R. sphenocephala in 1000-L outdoor experimental ponds. Three hybrid (F₁, backcrossj [B₁], backcross₂ [B₂]) and two parental (R. blairi [BB] and R. sphenocephala [SS]) larval genotypes were produced by artificial fertilzations using adult frogs from a natural population in central Missouri. Resultant larvae were reared in single-genotype populations and two-way mixtures at equal total numbers from hatching to metamorphosis. In single-genotype ponds, F₁ hybrid larvae had highest survival and BB were largest at metamorphosis. When F₁ and SS larvae were mixed together, F₁ hybrids had reduced survival and both F₁ and SS larvae metamorphosed at larger body masses than when reared separately. When mixed, both B₁ and SS larvae had shorter larval period lengths than when reared alone. Higher proportion of B₁ metamorphs were produced when larvae were mixed with either parental species than when reared alone. Larval fitness components as measured by survival, body mass at metamorphosis, proportion of survivors metamorphosing, and larval period length for B₂ hybrid and BB larvae were similar in single-genotype populations and mixtures. Comparison of composite fitness component estimates indicated hybrid genotypes possess equivalent or higher larval fitness relative to both parental species for the life-history fitness components measured. Despite reduced survival of F₁ hybrids in mixtures, backcross-generation hybrid genotypes demonstrated high levels of larval growth, survival, and metamorphosis in mixtures with parental species. Consequently, this study suggests natural hybridization and subsequent backcrossing between R. blairi and R. sphenocephala can produce novel and relatively fit hybrid genotypes capable of successful existence with parental species larvae. Thus, the evolutionary fate of hybrid and parental genotypes in this system may be influenced by exogenous selection mediated by genotypic composition of larval assemblages.     

Canopy Closure,Competition, and the Endangered Dusky Gopher Frog

Abstract: A major challenge facing wildlife biologists is understanding why some species go extinct while others persist in the same habitat. To address this question, we investigated whether tree canopy closure over ponds affects growth and survival of rare and common tadpoles within ponds and mediates competitive interactions among species. We conducted 2 experiments to test whether canopy closure and competition may have contributed to the decline of the endangered dusky gopher frog (Rana sevosa), but allowed the persistence of the southern leopard frog (R. sphenocephala). We explored the response of both species to canopy closure in single-species and mixed- (1:1) species treatments of identical total tadpole density. An experiment using aquatic enclosures in temporary ponds showed that canopy closure reduced tadpole growth approximately 20% for both species. Survival of dusky gopher frog tadpoles was higher in mixed-species enclosures than in single-species enclosures. In a complementary experiment using artificial ponds, dusky gopher frogs had lower survival to metamorphosis, reduced size at metamorphosis, and produced a lower total biomass of metamorphosed juveniles in shaded ponds. Southern leopard frogs exhibited reduced body size at metamorphosis only when shaded. These studies suggest that pond canopy closure, not larval competition, may be contributing to the decline of the dusky gopher frog. The different responses to canopy closure suggest a potential mechanism for the loss of dusky gopher frogs and the persistence of southern leopard frogs. Removal of trees from historically open-canopy ponds may help facilitate the recovery of dusky gopher frogs and benefit similar species.     

A comparative study of competitive ability between two cactophilic species in their natural hosts

Abstract Competition is a major aspect of the ecology of insect communities exploiting ephemeral and fragmented resources. We analysed the effect of intraspecific (single species culture) and interspecific (mixed species culture) competition on larval viability, developmental time and wing length in the cactophilic Drosophila buzzatii and Drosophila koepferae (Diptera: Drosophilidae) reared in cultured media prepared with fermenting tissues of three common natural cactus hosts in nature at different densities. Our results show that all traits measured were affected by both intra‐ and interspecifc competition, although the effect of competition depended on the Drosophila species and the rearing cactus. In fact, flies tended to have a lower viability, shorter wing size and longer developmental time as a function of increasing density in single species culture in both D. buzzatii and D. koepferae (intraespecific competition). Besides, the performance of both species was seriously affected (shorter body size, slower developmental times, lower viability) by the presence of heterospecific competitors except in the case of D. koepferae reared in its primary host plant, Trichocereus terschekii. We also show that D. koepferae successfully utilized Opuntia quimilo, which is absent in most parts of its distribution range. We discuss the roles of intra‐ and interspecific competition as determinants of the relative abundance of these two species in the arid zones of Southern South America.     

Influence of priority effects and pond location on invaded larval amphibian communities

In Florida, the Cuban Treefrog (Osteopilus septentrionalis) is a superb colonist and appears to be a significant driver of amphibian community dynamics. Decline of native anurans has been linked to possible competition with adult O. septentrionalis but interactions during the larval stage are largely unknown. Rearing O. septentrionalis tadpoles along with two native anurans, the Squirrel Treefrog (Hyla squirella) and the Southern Toad (Bufo terrestris) in both experimental artificial ponds and laboratory aquaria, the role of competition as the mechanism driving the dynamics of invaded amphibian communities in Florida was examined. Also examined was the role of priority effects and variation between pond locations in altering interactions between O. septentrionalis and native anuran larvae. Interspecific competition was strong during the larval stage; the presence of O. septentrionalis reduced larval performance and survival of native anurans. Pond location alone had little effect on interspecific interactions, but priority effects were strong. Pond location and priority effects acted together to influence species interactions. The selective influence of different interaction modifiers acted to increase or decrease the impacts of exotic species on native taxa.     

Complex competitive interactions in four species of Lucilia blowflies

Abstract.

• 1 The effects of intraspecific and interspecific larval competition on larval survival, adult size, adult longevity and fecundity were quantified in four species of coexisting Lucilia blowflies: illustris, silvarum, sericata and caesar.
• 2 There was a general negative effect of increasing density on larval and adult survival, size and fecundity. Additionally, complex species-specific and frequency-dependent responses were identified, which were not expected in these biologically and morphologically closely similar species.
• 3 Lucilia illustris, the numerically dominant species in the natural community, was a superior competitor to L.silvarum at intermediate densities but an inferior competitor at high density. Such nonlinear responses may be related to differences in the life histories and larval behaviour of the species (bigger eggs and more contest-type outcome of competition in L.silvarum).
• 4 We parameterized a model of interspecific competition on a subdivided resource in an attempt to reconcile the conflicting results on larval competitive abilities and the abundances of the species in the field. Using laboratory and field-estimated parameter values the model predicted coexistence of L.illustris and L.silvarum and the observed numerical dominance of the former species. The average densities of flies in the field are limited to relatively low levels, apparently preventing L.silvarum (the superior competitor at high density) from dominating and excluding L. illustris.

     

Amphibian lipid levels at metamorphosis correlate to post-metamorphic terrestrial survival

In organisms that have complex life cycles, factors in the larval environment may affect both larval and adult traits. For amphibians, the postmetamorphic transition from the aquatic environment to terrestrial habitat may be a period of high juvenile mortality. We hypothesized that lipid stores at metamorphosis may affect an animal’s success during this critical transition period. We examined variation in total lipid levels among years and sites in recently metamorphosed individuals of two pond-breeding salamander species, the marbled salamander (Ambystoma opacum) and the mole salamander (A. talpoideum), with limited data for one anuran species (southern leopard frog, Rana sphenocephala). Lipid levels were allometrically related to body size and ranged from 1.9 to 23.8% of body dry mass. The two salamander species differed in lipid allocation patterns, with A. opacum apportioning a higher percentage of total lipid reserves into fat bodies than A. talpoideum. Species differences in lipid allocation patterns may primarily reflect that large metamorphs will mature as one-year olds, and, regardless of species, will alter lipid compartmentalization accordingly. We used mark–recapture data obtained at drift fences encircling breeding ponds for 13 A. opacum cohorts to estimate the proportion of postmetamorphic individuals that survived to breed (age 1–4) and the mean age at first reproduction. Regression models indicated that size-corrected lipid level at metamorphosis (i.e., lipid residuals), and to a lesser extent rainfall following metamorphosis, was positively related to adult survival. Snout-vent length at metamorphosis was negatively related to age at first reproduction. We suggest that lipid stores at metamorphosis are vital to juvenile survival in the months following the transition from aquatic to terrestrial habitat, and that a trade-off shaped by postmetamorphic selection in the terrestrial habitat exists between allocation to energy stores versus structural growth in the larval environment.     

Caddisfly life histories along permanence gradients in high-altitude wetlands in Colorado (U.S.A.)

SUMMARY 1. Larvae of cased caddisflies (Limnephilidae and Phryganeidae) are among the most abundant and conspicuous invertebrates in northern wetlands. Although species replacements are often observed along permanence gradients, the underlying causal mechanisms are poorly understood. In this paper, we report on the distributional patterns of caddisflies in permanent and temporary high‐altitude ponds, and how those patterns reflect differences in life history characteristics that affect desiccation tolerance (fundamental niches) versus constraints related to biotic interactions (realised niches). 2. Species (Hesperophylax occidentalis and Agrypnia deflata) that were encountered only in permanent ponds are restricted in distribution by life history (no ovarian diapause, aquatic oviposition, and/or inability to tolerate desiccation). Although the egg masses of H. occidentalis tolerate desiccation, the larvae leave the protective gelatinous matrix of the egg mass because adults oviposit in water. 3. Three species (Asynarchus nigriculus, Limnephilus externus and L. picturatus) have life history characteristics (rapid larval growth, ovarian diapause and terrestrial oviposition of desiccation‐tolerant eggs) that should facilitate the use of both permanent and temporary habitats. However, A. nigriculus is rare or absent in most permanent ponds, and L. externus and L. picturatus are rare or absent in most temporary ponds. Experimental data from a previous study on the combined effects of salamander predation and interspecific interactions among caddisflies (e.g. intraguild predation) suggest that biotic interactions limit each species to a subset of potentially exploitable habitats. 4. Many wetland invertebrates exhibit species replacements along permanence gradients, but few studies have separated the relative importance of the effects of drying per se from the effects of biotic interactions. Our results emphasise the complementary roles of comparative data on life histories and experimental data on competition and predation for understanding invertebrate distributions along permanence gradients.     

Rhododendron aureum Georgi formed a special soil microbial community and competed with above‐ground plants on the tundra of the Changbai Mountain,China

Rhododendron aureum Georgi is a perennial evergreen dwarf shrub that grows at all elevations within the alpine tundra of northern China. Previous research has investigated the plant communities of R. aureum; however, little information is available regarding interspecific competition and underground soil microbial community composition. The objective of our study was to determine whether the presence of R. aureum creates a unique soil microbiome and to investigate the relationship between R. aureum and other plant species. Our study site ranged from 1,800 to 2,600 m above sea level on the northern slope of the Changbai Mountain. The results show that the soil from sites with an R. aureum community had a higher abundance of nitrogen‐fixing bacteria and a higher resistance to pathogens than soils from sites without R. aureum. We emphasize that R. aureum promotes a unique soil microbial community structure that is distinct from those associated with other plants. Elevation and microbial biomass were the main influencing factors for plant community structure. Analysis of interspecific relationships reveals that R. aureum is negatively associated with most other dominant shrubs and herbs, suggesting interspecific competition. It is necessary to focus on other dominant species if protection and restoration of the R. aureum competition is to occur. In the future, more is needed to prove whether R. aureum decreases species diversity in the tundra ecosystems of Changbai Mountain.     

Coexistence between two fruit fly species is supported by the different strength of intra‐ and interspecific competition

1. Unravelling the strength and modes of interspecific interactions between resident and introduced species is necessary in order to understand the basis of their coexistence or the displacement of the former by the latter. In Argentina, the indigenous Tephritidae fly Anastrepha fraterculus overlaps its distribution and host fruit with the introduced species Ceratitis capitata. 2. This study focused on the relative strength of intra‐ and interspecific competition during the larval stage as a potential factor supporting coexistence. Classical competition experiments (addition and substitution) were conducted between larvae of the two species reared in artificial larval diet. The study evaluated whether a temporal separation between oviposition events affects the outcome of the competition. 3. When both species started to consume the resource at the same time, A. fraterculus experienced a negative effect in larval survival, pupal weight and duration of larval stage, while for C. capitata, pupal weight decreased. When A. fraterculus started feeding 1 day earlier than C. capitata, the negative effects became milder, and when the temporal separation increased, these effects were reversed. Substitution experiments showed an increase in pupal weight when larvae had to share the resource with heterospecific larvae, and showed negative effects suffered for both species when they shared the resource with conspecific individuals. 4. These results suggest that intraspecific competition is stronger than interspecific competition, and a differential oviposition preference could generate an asynchrony of these species in nature. Such mechanisms could favour coexistence between A. fraterculus and C. capitata in an environment previously occupied only by the former.     

Competition between Bufo larvae in a eutrophic pond

Competition between larvae of two anuran species (Bufo bufo and B. calamita) was investigated under field conditions likely to disfavour cell-mediated interference mechanisms. The experiment used triplicated cage treatments in an unshaded farm pond, a poor habitat for the unicellular pathogen Anurofeca richardsi implicated in interference competition between these anurans in sand dune ponds. The farm pond experienced lower maximum temperatures than a nearby dune pond but sustained larger numbers of eukaryotic algae and therefore had higher primary productivity. Survival and growth of B. calamita larvae were inversely related to density in all treatments but interspecific effects were much more severe than intraspecific ones. There was no evidence of A. richardsi in any treatment and competition between the Bufo larvae was therefore intense in the absence of Anurofeca-mediated interference effects. Anuran larvae reduced the standing crop and altered the community composition of algae in the treatment cages but larval growth rates were not simply related to food availability. Algal cell numbers in larval guts, a measure of food acquisition, were however inversely related to tadpole density in both species. Feeding niche overlap was high but decreased as larval density increased. Resource competition was implicated as the most probable major mechanism. Received: 1 September 1999 / Accepted: 20 January 2000     

Larval consumption rates,interspecific predation,and local guild composition of egg-overwintering Agabus (Coleoptera,Dytiscidae) species in vernal ponds

Summary In vernal ponds in the boreal region, egg-over-wintering Agabus species form a guild that feeds mainly on larvae and pupae of aedine mosquitoes. The regular co-existence of very similar Agabus species indicates local communities not structured by interspecific competition. However, the lower number of species in local guilds than in the regional species pool poses a problem of limited membership. We suggest that the species of this guild display habitat differences mainly with respect to water temperature, pond size and prey density. In this view, habitat selection reflects body size and thermal growth response of the species, mainly in connection with larval development. We present field data from two northern Swedish vernal ponds. Based on these data, feeding experiments were performed to test the hypothesis outlined above. At a high prey density, larvae of all instars of the larger species A. erichsoni Gemm. & Har. had a significantly higher consumption rate than those of the smaller species A. opacus Aubé. At a low prey density the differences were smaller, and only the third instar larvae differed significantly. At 2° C, larvae of A. opacus had a significantly higher consumption rate than those of A. congener (Thunberg). At 15° C, no significant difference was observed. In studies of within-guild interspecific predation, always the larger larvae consumed the smaller ones. Field data show that egg hatching is spread out in time, and show interspecific differences. Consequently, the effects of unexpected droughts differ with species.     

Interspecific competition and coexistence of the two chrysomelids,Gastrophysa atrocyanea motschulsky andGalerucella vittaticollis baly (Coleoptera: Chrysomelidae), under limited food resource conditions

Field experiments were conducted in order to investigate the mode of exploitation of food resources and the mechanism of coexistence of mixed larval populations of the two chrysomelids,Gastrophysa atrocyanea andGalerucella vittaticollis, under limited food resource conditions. The larval survival rates seemed high enough to assure coexistence when hatchlings of the two species were released in 1∶1 and 1∶3 ratios on a host plant. However, the survival rate became almost nil for both species when a 3∶1 ratio was employed, suggesting asymmetrical interspecific competition. Wasted food consumption was much higher inG. atrocyanea larvae. The population ofG. atrocyanea seemed to be regulated more by intraspecific competition, while on the other hand, the population ofG. vittaticollis was considered to be more likely affected by the interspecific competition withG. atrocyanea, depending on the initial ratio and density of the two species.     

Vegetative composition and community structure associated with beaver ponds in Canaan valley, West Virginia, USA

Beavers (Castor canadensis) can cause dramatic changes in vegetative composition and diversity. Although alterations by beaver have been studied extensively, little attention has been paid to the effects beaver impoundments have on rare plants. Effective conservation of riparian and wetland rare plant species must consider the responses of vegetation to changes in hydrology that can occur when beaver populations are present. The goal of this research was to establish the occurrence of locally rare plant species, examine community composition, and analyze vegetative community structure of vegetation associated with beaver ponds in Canaan Valley, West Virginia, USA. Species richness and diversity were similar between plots located inside beaver ponds and adjacent to beaver ponds (P > 0.05). Although no significant difference in rare plant species was detected among pond ages, the oldest ponds (>56 years) had twice as many rare species as the youngest ponds (≤6 years). The youngest ponds had higher overall mean species richness (S) than ponds 7–56 years old (P < 0.05), but S returned to similar levels in the oldest ponds. Of the 15 rare species observed, most were classified as obligate (9) or facultative wetland (4) species. The youngest ponds contained the fewest number of rare species. Multiple response permutation procedure (MRPP) analyses of community structure detected no relation between community composition and either pond age or size. However, both MRPP and non-metric multi-dimensional scaling showed proximity to pond was important in herbaceous community structure. Wetlands in beaver ponds also were shown to be distinct from adjacent wetland areas. Conservation of existing beaver populations is necessary so that the entire spectrum of pond ages is available for the maintenance of rare plant species and communities.     

Differentiated egg size of the cannibalistic salamander <Emphasis Type="Italic">Hynobius retardatus</Emphasis>

Larvae of the salamander, Hynobius retardatus, are carnivorous, and even though there are two morphs, a typical morph and a broad-headed or “cannibal” morph, both are cannibalistic. They also sometimes eat other large prey, for example larvae of the frog, Rana pirica. In natural habitats, use of both conspecific and R. pirica larvae as food may contribute more strongly to high survival and substantially to fitness when larval densities are higher, because early-stage H. retardatus larvae sometimes experience scarcity of their typical prey. In cannibalistic oviparous amphibians, larger individuals that developed from larger eggs can more efficiently catch and consume larger prey and thus their survival may be better than that of smaller individuals developed from smaller eggs. Populations might therefore diverge in respect of egg size in response to variation in the density of conspecific and R. pirica larvae in natural ponds, with eggs being larger when larval density is higher. I examined how variance in hatchling size correlated with the incidence of cannibalism, and whether increasing larval density in natural ponds correlated with increasing egg size. Variance in initial larval body size facilitated cannibalism, and egg size increased as larval density in the ponds increased. In ponds with high larval density, where cannibalism and large prey consumption is a critical factor in offspring fitness, the production of fewer clutches with larger eggs, and thus of fewer and larger offspring, results in greater maternal fitness. Variation among the mean egg size in populations is likely to represent a shift in optimum egg size across larval density gradients.     

Density of an intraguild predator mediates feeding group size, intraguild egg predation, and intra- and interspecific competition

Intraguild predation (IGP) is common in most communities, but many aspects of density-dependent interactions of IG predators with IG prey are poorly resolved. Here, we examine how the density of an IG predator can affect feeding group size, IG egg predation, and the growth responses of IG prey. We used laboratory feeding trials and outdoor mesocosm experiments to study interactions between a social intraguild predator (larvae of the wood frog; Rana sylvatica) and its prey (spotted salamander; Ambystoma maculatum). Larvae of R. sylvatica could potentially affect A. maculatum by consuming shared larval food resources or by consuming eggs and hatchlings. However, successful egg predation requires group feeding by schooling tadpoles. We established from five to 1,190 hatchlings of R. sylvatica in mesocosms, then added either 20 A. maculatum hatchlings to study interspecific competition, or a single egg mass to examine IGP. Crowding strongly suppressed the growth of R. sylvatica, and IGP was restricted to the egg stage. In the larval competition experiment, growth of A. maculatum was inversely proportional to R. sylvatica density. In the predation experiment, embryonic mortality of A. maculatum was directly proportional to the initial density of R. sylvatica and the mean number of tadpoles foraging on egg masses. IGP on eggs reduced A. maculatum hatchling density, which accelerated larval growth. Surprisingly, the density of R. sylvatica had no overall effect on A. maculatum growth because release from intraspecific competition via egg predation was balanced by increased interspecific competition. Our results demonstrate that the density of a social IG predator can strongly influence the nature and intensity of interactions with a second guild member by simultaneously altering the intensity of IGP and intra- and interspecific competition.L . A. Burley and A. T. Moyer contributed equally to this work.     

EVOLUTION OF GIGANTISM IN NINE‐SPINED STICKLEBACKS

The relaxation of predation and interspecific competition are hypothesized to allow evolution toward “optimal” body size in island environments, resulting in the gigantism of small organisms. We tested this hypothesis by studying a small teleost (nine‐spined stickleback, Pungitius pungitius) from four marine and five lake (diverse fish community) and nine pond (impoverished fish community) populations. In line with theory, pond fish tended to be larger than their marine or lake conspecifics, sometimes reaching giant sizes. In two geographically independent cases when predatory fish had been introduced into ponds, fish were smaller than those in nearby ponds lacking predators. Pond fish were also smaller when found in sympatry with three‐spined stickleback (Gasterosteus aculeatus) than those in ponds lacking competitors. Size‐at‐age analyses demonstrated that larger size in ponds was achieved by both increased growth rates and extended longevity of pond fish. Results from a common garden experiment indicate that the growth differences had a genetic basis: pond fish developed two to three times higher body mass than marine fish during 36 weeks of growth under similar conditions. Hence, reduced risk of predation and interspecific competition appear to be chief forces driving insular body size evolution toward gigantism.     

Competition between invasive and indigenous species: an insular case study of subterranean termites

An important requirement for the management of invasive species is to identify the biological and ecological factors that influence the ability of such species to become established and spread within a new environment. Although competition is one of the key interactions determining the coexistence of species and exclusion, few studies directly examine the mechanism of competitive interactions within invasive communities. This study focused on putative competition in a social insect invader, R. flavipes, an American termite introduced into France, and an indigenous European termite, R. grassei. We first characterized and mapped a zone of sympatry between these two species. We then evaluated the degree of direct and indirect competition by comparing several life-history traits: behavioral aggression, chemical recognition and dispersion modes. Interspecific competition revealed that R. flavipes was dominant over R. grassei. Intraspecific competition was not found in R. flavipes while it appeared in varying degrees in R. grassei. These findings seemed to be correlated with the remarkable chemical homogeneity found in R. flavipes in comparison with R. grassei. Genetic analyses revealed that R. flavipes foraged over a greater distance than R. grassei colonies and might suggest a difference in the capacity to produce secondary reproductives. These findings suggest that R. flavipes has a significant advantage owing to competitive asymmetry that may enable the species to become dominant. The interspecific superiority, lack of intraspecific aggression and large extensive colonies, seem to be some of the reasons for its invasive success.     

Competition in slow motion: the unusual case of benthic marine communities in the wake of the end‐Permian mass extinction

Changes of community structure in response to competition usually take place on timescales that are much too short to be visible in the geological record. Here we report the notable exception of a benthic marine community in the wake of the end‐Permian mass extinction, which is associated with the microbial limestone facies of the earliest Triassic of South China. The newly reported fauna is well preserved and extraordinarily rich (30 benthic macroinvertebrate species, including the new species Astartella? stefaniae (Bivalvia) and Eucochlis obliquecostata (Gastropoda)) and stems from an environmentally stable setting providing favourable conditions for benthic organisms. Whereas changes in the taxonomic composition are negligible over the observed time interval of 10–100 ka, three ecological stages are identified, in which relative abundances of initially rare species continuously increased at the cost of previously dominant species. Concomitant with the changes of dominant species is an increase in faunal evenness and heterogeneity. In the absence of both environmental and taxonomic changes, we attribute this pattern to the long‐term effects of interspecific competition, which acted at an unusually slow pace because the number of competing species and potential immigrants was dramatically reduced by the end‐Permian mass extinction. We suggest that these non‐actualistic conditions led to decreased rates of niche differentiation and hence to the delayed rediversification of benthos that characterizes the aftermath of the greatest Phanerozoic mass extinction event. A hyperbolic diversification model is proposed, which accounts for the positive relationship between the intensity of interspecific competition and the rate of niche differentiation and resolves the conundrum of delayed rediversification at a time when niche space was largely vacated.     

Variability of intra- and interspecific competitions of bamboo stump mosquito larvae over small and large spatial scales

We carried out field experiments to examine the variability of interspecific competition of mosquito larvae among microcosms in a bamboo grove (small spatial scale) and between bamboo groves at two sites, with single and multiple mosquito species (large spatial scale). Four types of microcosms that differed in capacity and litter input were set. In the hillside bamboo grove, where multiple species occurred, succession of the predominant species from Aedes albopictus to Tripteroides bambusa was observed in control microcosms from which no mosquito larvae were removed. Weekly removal of competitive species resulted in increased pupation of A. albopictus and adult body weight under both rich and poor resource conditions. In the late period of the experiments, the effect of competitor removal on pupation of A. albopictus was greater in deep containers that never dried than in shallow containers that were dried in the laboratory. The number of eggs showed a slight difference between competitor‐excluded and deep control microcosms. These results indicate that interspecific competition limits pupation of A. albopictus more strongly in deep containers than in shallow and drought‐prone containers.
Compared with the hillside site, the larval density of A. albopictus attained a higher density in the bamboo grove in the plain where no competitive species occurred, due to a higher oviposition rate. Lower rate of pupation and lower adult weight at the plain site than at the hillside site indicated that resource limitation was more severe at the plain site. Populations of A. albopictus at hillside and plain sites appeared to suffer from strong inter‐ and intraspecific competition, respectively. At the hillside site, the intensity of interspecific competition appeared to increase later in the breeding season, with a high larval density of T. bambusa. In contrast, at the plain site, intensity of intraspecific competition appeared to be reduced later in the breeding season with decreasing larval density of A. albopictus.