Reproductive interference via interspecific pairing in an amphipod species complex

Many species experience reproductive interference defined as interspecific mating interactions that reduce the fitness of individuals in at least one of the species. We examined reproductive interference among three cryptic amphipod species that co-occur in the vegetated zone of lakes. Because these animals form precopulatory pairs and males commonly have indiscriminant mating behavior, we predicted that (1) there would be interspecific pairing, (2) interspecific pairing would be more common when conspecifics were rare, and (3) interspecific pairing would be more common in species combinations where males are larger than females. Using color phenotypes to distinguish species, a novel discovery reported here, we conducted experiments to evaluate patterns of interspecific pairing. In a no-choice experiment (i.e., amphipods had access to mates of only one species), we observed a low rate (5 %) of interspecific pairing and 80 % of these cases involved males that were larger than females. In a second experiment where individuals had access to conspecific mates at varying relative abundance, only 0.04 % of the pairs observed were between heterospecifics, suggesting that the interspecific pairing observed in the no-choice experiment was an artifact of not having access to conspecific mates. Our results suggest that at least one sex has sufficient species-recognition abilities to circumvent the formation of interspecific pairs; therefore, reproductive interference may be minimal in these species.     

A classification framework for interspecific trade-offs in aquatic ecology

In some cases, wildlife management objectives directed at multiple species can conflict with one another, creating species trade-offs. For managers to effectively identify trade-offs and avoid their undesirable outcomes, they must understand the agents involved and their corresponding interactions. A literature review of interspecific trade-offs within freshwater and marine ecosystems was conducted to illustrate the scope of potential interspecific trade-offs that may occur. We identified common pitfalls that lead to failed recognition of interspecific trade-offs, including, single-species management and limited consideration of the spatial and temporal scale of ecosystems and their management regimes. We devised a classification framework of common interspecific trade-offs within aquatic systems. The classification can help managers determine whether the conflict is species based through direct relationships (i.e., predator–prey, competition, other antagonistic relationships) or indirect relationships involving intermediate species (i.e., conflict-generating species) or whether the conflict is driven by opposing management objectives for species that would otherwise not interact (i.e., nontarget management effects). Once the nature and scope of trade-offs are understood, existing decision-making tools, such as structured decision-making and real-options analysis, can be incorporated to improve the management of aquatic ecosystems. Article Impact Statement: A synthesis of interspecific trade-offs in aquatic ecosystems supports their identification and resolution.     

Salinity effects on the coexistence of cryptic species: a case study on marine nematodes

The coexistence of four cryptic species of Rhabditis (Pellioditis) marina (Nematoda: Rhabditidae) at small geographical scale challenges ecological competition theory and was therefore studied in the laboratory at two different salinities, where their performance in combined cultures was compared with that in monospecies cultures. We found that three of the four cryptic species were able to coexist, but that interspecific interactions (competition and facilitation) were common. Salinity had an effect on these interactions, with a shift from contest to scramble competition. This shift may result from an increased population development of two of the four species at the lower salinity in the monospecific cultures. This experiment demonstrates that abiotic conditions may play an important role in achieving coexistence between cryptic species and can alter the interspecific interactions between them.     

Invasional interference due to similar inter- and intraspecific competition between invaders may affect management

As the number of biological invasions increases, the potential for invader-invader interactions also rises. The effect of multiple invaders can be superadditive (invasional meltdown), additive, or subadditive (invasional interference); which of these situations occurs has critical implications for prioritization of management efforts. Carduus nutans and C. acanthoides, two congeneric invasive weeds, have a striking, segregated distribution in central Pennsylvania, U.S.A. Possible hypotheses for this pattern include invasion history and chance, direct competition, or negative interactions mediated by other species, such as shared pollinators. To explore the role of resource competition in generating this pattern, we conducted three related experiments using a response-surface design throughout the life cycles of two cohorts. Although these species have similar niche requirements, we found no differential response to competition between conspecifics vs. congeners. The response to combined density was relatively weak for both species. While direct competitive interactions do not explain the segregated distributional patterns of these two species, we predict that invasions of either species singly, or both species together, would have similar impacts. When prioritizing which areas to target to prevent the spread of one of the species, it is better to focus on areas as yet unaffected by its congener; where the congener is already present, invasional interference makes it unlikely that the net effect will change.     

Functional responses: a question of alternative prey and predator density

Throughout the study of ecology, there has been a growing realization that indirect effects among species cause complexity in food webs. Understanding and predicting the behavior of ecosystems consequently depends on our ability to identify indirect effects and their mechanisms. The present study experimentally investigates indirect interactions arising between two prey species that share a common predator. In a natural field experiment, we introduced different densities of mealworms (Tenebrio molitor), an alternative prey, to a previously studied predator-prey system in which paper wasps (Polistes dominulus) preyed on shield beetle larvae (Cassida rubiginosa). We tested if alternative prey affects predation on the first prey (i.e., the predator-dependent functional response of paper wasps) by modifying either interference among predators or the effective number of predators foraging on shield beetles. Presence of mealworms significantly reduced the effective number of predators, whereas predator interference was not affected. In this way, the experimentally introduced alternative prey altered the wasps' functional response and thereby indirectly influenced C. rubiginosa density. In all prey-density combinations offered, paper wasps constantly preferred T. molitor. This led to an asymmetrical, indirect interaction between both prey species: an increase in mealworm density significantly relaxed predation on C. rubiginosa, whereas an increase in C. rubiginosa density intensified predation on mealworms. Such asymmetrical outcomes of a fixed food preference can significantly affect the population dynamics of the species involved. In spite of the repeated finding of a Type III functional response in this system, our experiment did not reveal switching behavior in paper wasps. The variety of mechanisms underlying direct and indirect interactions within our study system exemplifies the importance of incorporating alternative prey when investigating the impact of a generalist predator on a focal prey population under realistic field conditions.     

Response of American redstarts (suborder Passeri) and least flycatchers (suborder Tyranni) to heterospecific playback: the role of song in aggressive interactions and interference competition

Least flycatchers (Empidonax minimus) and American redstarts (Setophaga ruticilla) overlap in the use of food resources on their breeding grounds, promoting high levels of interspecific aggression by the socially dominant flycatcher. We examined the role of song in this interspecific aggression by using repeated-measures-designed playback experiments and observational data on induced aggressive interactions. Flycatchers were more likely to approach the speaker during presentation of redstart song than during intervals of no song or presentation of control song. Approach was close enough to enable visual contact with a singing redstart. In contrast, redstarts made significantly fewer flights following presentation of flycatcher song, when risk of flycatcher attack may be greatest. Reducing the number of flights likely reduces the risk of flycatcher attack on the redstart, as flycatchers do not attack stationary redstart models and are apparently dependent on cues from redstart flight for visual heterospecific recognition. Flycatcher-specific responses of redstarts and marked differences in song morphology rule out misdirected intraspecific aggression as a proximate or ultimate cause of interspecific response to song. Results indicate that song is an important component in aggressive interactions between these two species, and reflect the dominant role of the flycatcher in such interactions. Our results also illustrate the capacity for interspecific interference competition to influence behavior and heterospecific song recognition in two distant avian taxa. Received: 10 January 1995/Accepted after revision: 8 June 1996     

Coexistence of three specialist aphids on common milkweed, Asclepias syriaca

Coexistence of host-specific herbivores on plants is believed to be governed by interspecific interactions, but few empirical studies have systematically unraveled these dynamics. We investigated the role of several factors in promoting coexistence among the aphids Aphis nerii, Aphis asclepiadis, and Myzocallis asclepiadis that all specialize on common milkweed (Asclepias syriaca). Competitive exclusion is thought to occur when interspecific competition is stronger than intraspecific competition. Consequently, we investigated whether predators, mutualists, or resource quality affected the strength of intra- vs. interspecific competition among aphids in factorial manipulations of competition with exposure to predation, ants, and variable plant genotypes in three separate experiments. In the predation x competition experiment, predators reduced aphid per capita growth by 66%, but the strength of intra- and interspecific competition did not depend on predators. In the ants x competition experiment, ants reduced per capita growth of A. nerii and M. asclepiadis (neither of which were mutualists with ants) by approximately one-half. In so doing, ants ameliorated the negative effects of these competitors on ant-tended A. asclepiadis by two-thirds, representing a novel benefit of ant-aphid mutualism. Nevertheless, ants alone did not explain the persistence of competitively inferior A. asclepiadis as, even in the presence of ants, interspecific competition remained stronger than intraspecific competition. In the plant genotype x competition experiment, both A. asclepiadis and M. asclepiadis were competitively inferior to A. nerii, with the strength of interspecific competition exceeding that of intraspecific competition by 83% and 23%, respectively. Yet these effects differed among milkweed genotypes, and there were one or more plant genotypes for each aphid species where coexistence was predicted. A synthesis of our results shows that predators play little or no role in preferentially suppressing competitively dominant A. nerii. Nonetheless, A. asclepiadis benefits from ants, and A. asclepiadis and M. asclepiadis may escape competitive exclusion by A. nerii on select milkweed genotypes. Taken as a whole, the coexistence of three host-specific aphid species sharing the same resource was promoted by the dual action of ants as antagonists and mutualists and by genetic diversity in the plant population itself.     

Hybridization and the Extinction of Rare Plant Species

Much has been written about the role of interspecific competition, disease, herbivory, and the loss of key mutualisms in the extinction of rare plant species. Interspecific hybridization rarely is considered among the biotic interactions that promote extinction. We show how hybridization may contribute to the demise of rare plant species through demographic swamping and genetic assimilation by an abundant congener. We contend that the growth of the hybrid subpopulation is the key to rare species assimilation, and we show how the production of hybrid seed, the fitness of hybrids, and pest pressure affect hybrid proliferation. We also discuss how habitat disturbance, unspecialized pollinators, and weak crossing barriers promote hybridization, and how the negative consequences of hybridization are unlikely to be compensated for by immigration from conspecific populations. We also illustrate stages in the demise of species in island floras. We suggest that hybridization is an increasing threat to rare species because ecological barriers are being disrupted by human activities.     

Variability of competition at scales of 10<Superscript>1</Superscript>, 10<Superscript>3</Superscript>, 10<Superscript>5</Superscript>, and 10<Superscript>6</Superscript> m: encrusting arctic community patterns

Variability in interference competition was studied in benthic marine communities of the arctic and subarctic Atlantic intertidal and shallow subtidal zones. We sampled multiple square-metre quadrats at distances of 10¹, 10³, and 10⁵ m apart around the high polar island of Spitsbergen (Svalbard Archipelago). We also took some similar samples in Iceland and in the Faeroe Islands (10⁶ m apart from Spitsbergen samples). Encrusting fauna were present on high arctic intertidal rocks but we only found competitive interactions on subtidal substrata. On subarctic Icelandic and Faeroese shores, in contrast, spatial competition was common even in the intertidal zone. Analysis of variance of competition intensity data (numbers of interactions per area) revealed multiple factors to be significant influences explaining variability. Amongst the 10¹-, 10³-, and 10⁵-m spatial scales, only the largest emerged as a significant term. Whether intra- or interspecific competition dominated the types of interactions varied greatly between sites: 21–97% of competition was intraspecific. The proportion of competitive encounters resulting in a decided outcome (i.e. a win for one competitor and a loss for the other, rather than a tie or standoff between them) showed little variability at any scale. All the values of competition transitivity (how hierarchical a pecking order is) were very high compared to values reported in the literature from any other (polar or non-polar) locality. Variability in this measure was generally <10% across scales. We conclude from our data that great care must be taken in interpreting patterns of competition between similar taxa in large-scale space or time. Not only did most aspects of competition in our study communities vary significantly at the 10⁵-m scale but different aspects of competition varied at different scales and by hugely different amounts.Communicated by J.P. Thorpe, Port Erin     

An experimental study of competition between fire ants and Argentine ants in their native range

An understanding of why introduced species achieve ecological success in novel environments often requires information about the factors that limit the abundance of these taxa in their native ranges. Although numerous recent studies have evaluated the importance of natural enemies in this context, relatively few have examined how ecological success may result from differences in the magnitude of interference competition between communities in the native and introduced ranges of nonnative species. Here we examine how native-range competitive environments may relate to invasion success for two important invasive species, the red imported fire ant (Solenopsis invicta) and the Argentine ant (Linepithema humile), in a region of native-range sympatry. At two study sites in northern Argentina, we used stable-isotope analysis, a variety of observational approaches, and two different reciprocal removal experiments to test (1) whether S. invicta competes asymmetrically with L. humile (as suggested by the 20th century pattern of replacement in the southeastern United States) and (2) the extent to which these two species achieve behavioral and numerical dominance. Stable-isotope analysis and activity surveys indicated that S. invicta and L. humile are both omnivores and forage during broadly overlapping portions of the diel cycle. Short-term removal experiments at baits revealed no competitive asymmetry between S. invicta and L. humile. Longer-term colony removal experiments illustrated that S. invicta and L. humile experience an approximately equal competitive release upon removal of the other. Our results indicate that neither S. invicta nor L. humile achieves the same degree of behavioral or ecological dominance where they co-occur in native populations as they do in areas where either is common in their introduced range. These results strongly suggest that interspecific competition is an important limiting factor for both S. invicta and L. humile in South America.     

Neighborhood analyses of canopy tree competition along environmental gradients in New England forests.

We use permanent-plot data from the USDA Forest Service's Forest Inventory and Analysis (FIA) program for an analysis of the effects of competition on tree growth along environmental gradients for the 14 most abundant tree species in forests of northern New England, USA. Our analysis estimates actual growth for each individual tree of a given species as a function of average potential diameter growth modified by three sets of scalars that quantify the effects on growth of (1) initial target tree size (dbh), (2) local environmental conditions, and (3) crowding by neighboring trees. Potential growth of seven of the 14 species varied along at least one of the two environmental axes identified by an ordination of relative abundance of species in plots. The relative abundances of a number of species were significantly displaced from sites where they showed maximum potential growth. In all of these cases, abundance was displaced to the more resource-poor end of the environmental gradient (either low fertility or low moisture). The pattern was most pronounced among early successional species, whereas late-successional species reached their greatest abundance on sites where they also showed the highest growth in the absence of competition. The analysis also provides empirical estimates of the strength of intraspecific and interspecific competitive effects of neighbors. For all but one of the species, our results led us to reject the hypothesis that all species of competitors have equivalent effects on a target species. Most of the individual pairwise interactions were strongly asymmetric. There was a clear competitive hierarchy among the four most shade-tolerant species, and a separate competitive hierarchy among the shade-intolerant species. Our results suggest that timber yield following selective logging will vary dramatically depending on the configuration of the residual canopy, because of interspecific variation in the magnitude of both the competitive effects of different species of neighbors and the competitive responses of different species of target trees to neighbors. The matrix of competition coefficients suggests that there may be clear benefits in managing for specific mixtures of species within local neighborhoods within stands.     

Coupling stable isotopes with bioenergetics to estimate interspecific interactions.

Interspecific interactions are often difficult to elucidate, particularly with large vertebrates at large spatial scales. Here, we describe a methodology for estimating interspecific interactions by combining stable isotopes with bioenergetics. We illustrate this approach by modeling the population dynamics and species interactions of a suite of vertebrates on Santa Cruz Island, California, USA: two endemic carnivores (the island fox and island spotted skunk), an exotic herbivore (the feral pig), and their shared predator, the Golden Eagle. Sensitivity analyses suggest that our parameter estimates are robust, and natural history observations suggest that our overall approach captures the species interactions in this vertebrate community. Nonetheless, several factors provide challenges to using isotopes to infer species interactions. Knowledge regarding species-specific isotopic fractionation and diet breadth is often lacking, necessitating detailed laboratory studies and natural history information. However, when coupled with other approaches, including bioenergetics, mechanistic models, and natural history, stable isotopes can be powerful tools in illuminating interspecific interactions and community dynamics.     

Intratree vertical variation of fruit density and the nature of contest competition in frugivores

Recently, we demonstrated that the highest densities of fruit pulp are located in the uppermost zones of tree crowns. Since heterogeneous distributions of depletable food is theorized to foster contest competition, we tested three hypotheses involving rank differences among species of arboreal frugivores: (1) In the absence of competitors, species tend to feed in higher strata of tree crowns; (2) interspecific contest competition occurs through monopolization and usurpation of feeding sites in these higher strata; and (3) subordinate species decrease their feeding height and ingestion rate when dominants enter the food patch. To test these hypotheses, we observed chimpanzees (Pan troglodytes), red-tailed monkeys (Cercopithecus ascanius), blue monkeys (Cercopithecus mitis), and gray-cheeked mangabeys (Lophocebus albigena) in Kibale National Park, Uganda. We found that: (1) all four primates fed preferentially in upper tree crowns when alone, (2) dominant species monopolized and aggressively usurped the upper crown when co-feeding with subordinates and the latter retreated below the middle of tree crowns, (3) in the presence of dominant species, subordinate species showed lower standardized feeding height and modified their food intake rates, while dominants were not affected by the subordinate species, (4) subordinates moved down at the arrival of and up at the departure of dominants, and (5) the presence of folivores in the tree did not affect the feeding height of a frugivore, even through folivores were socially dominant. Contrary to expectations, we found that red-tailed monkeys decreased their movements between successive fruits that they ate in the presence of blue monkeys compared to when they were feeding alone, perhaps to avoid disturbing dominants and attracting aggression or because they ingested more semi-ripe and green unripe fruits, i.e., more food of lower quality.     

Increasing enemy biodiversity strengthens herbivore suppression on two plant species

Concern over biodiversity loss, especially at higher trophic levels, has led to a surge in studies investigating how changes in natural enemy diversity affect community and ecosystem functioning. These studies have found that increasing enemy diversity can strengthen, weaken, and not affect prey suppression, demonstrating that multi-enemy effects on prey are context-dependent. Here we ask how one factor, plant species identity, influences multi-enemy effects on prey. We focused on two plant species of agricultural importance, potato (Solanum tuberosum), and collards (Brassica oleracea L.). These species share a common herbivorous pest, the green peach aphid (Myzus persicae), but vary in structural and chemical traits that affect aphid reproductive rates and which may also influence inter-enemy interactions. In a large-scale field experiment, overall prey exploitation varied dramatically among the plant species, with enemies reducing aphid populations by approximately 94% on potatoes and approximately 62% on collards. Increasing enemy diversity similarly strengthened aphid suppression on both plants, however, and there was no evidence that plant species identity significantly altered the relationship between enemy diversity and prey suppression. Microcosm experiments suggested that, on both collards and potatoes, intraspecific competition among natural enemies exceeded interspecific competition. Enemy species showed consistent and significant differences in where they foraged on the plants, and enemies in the low-diversity treatment tended to spend less time foraging than enemies in the high-diversity treatment. These data suggest that increasing enemy diversity may strengthen aphid suppression because interspecific differences in where enemies forage on the plant allow for greater resource partitioning. Further, these functional benefits of diversity appear to be robust to changes in plant species identity.     

Rules of engagement for adult salamanders in territorial conflicts with heterospecific juveniles

Coexisting species within a guild have the potential for resource overlap and consequently for interspecific competition (e.g., interspecific territoriality). When the adults are of different sizes, which frequently is the case in terrestrial salamander communities in eastern North America, competition may occur between juveniles of the larger species and adults of the smaller species. Adults of the relatively small redbacked salamander (Plethodon einereus: up to 13 cm total length) defend intra- and intersexual territories on the forest floor, and they are broadly sympatric with the larger P. glutinosus (up to 21 cm total length). Although individuals of the two species share the same forest floor habitat, we found significantly fewer juveniles of P. glutinosus sharing territories with 336 same-size adults of P. cinereus than would be expected by random chance alone. In laboratory experiments, residential adults of P. cinereus were as aggressive toward juvenile intruders of P. glutinosus as they were toward adult conspecific intruders. Therefore, adults of P. cinereus appear to defend territories against juveniles of P. glutinosus, illustrating how interference competition may depend on the symmetry of sizes between the species.     

The invasibility of marine algal assemblages: role of functional diversity and identity

The emergence of the biodiversity-ecosystem functioning debate in the last decade has renewed interest in understanding why some communities are more easily invaded than others and how the impact of invasion on recipient communities and ecosystems varies. To date most of the research on invasibility has focused on taxonomic diversity, i.e., species richness. However, functional diversity of the communities should be more relevant for the resistance of the community to invasions, as the extent of functional differences among the species in an assemblage is a major determinant of ecosystem processes. Although coastal marine habitats are among the most heavily invaded ecosystems, studies on community invasibility and vulnerability in these habitats are scarce. We carried out a manipulative field experiment in tide pools of the rocky intertidal to test the hypothesis that increasing functional richness reduces the susceptibility of macroalgal communities to invasion. We selected a priori four functional groups on the basis of previous knowledge of local species characteristics: encrusting, turf, subcanopy, and canopy species. Synthetic assemblages containing one, two, three, or four different functional groups of seaweeds were created, and invasion by native species was monitored over an eight-month period. Cover and resource availability in the assemblages with only one functional group showed different patterns in the use of space and light, confirming true functional differences among our groups. Experimental results showed that the identity of functional groups was more important than functional richness in determining the ability of macroalgal communities to resist invasion and that resistance to invasion was resource-mediated.     

杉木-观光木人工混交林竞争及邻体干扰指数研究

将杉木和观光木人工混交林中林木竞争影响空间划分为内、外两圈,在此基础上将竞争指数分解为不同的邻体干扰指数,对象木生长量与3种干扰指数的回归分析表明,以杉木为对象木时,利用张大勇提出的干扰指数模型(Izdy)效果最好;以观光木为对象木时,利用Weiner提出的干扰指数模型(Iwg)效果最好.同时,竞争强度研究表明,杉木种内竞争>杉木观光种间竞争>观光木种内竞争.通过对几种竞争影响的划分比较,说明将竞争空间分为内、外两圈能较好地反应林木的竞争规律.表3参17     

Spatial and behavioral interactions between a native and introduced salamander species

Behavioral interactions with native species may influence the invasiveness of introduced species. The salamanders Plethodon glutinosus and P. jordani in the eastern United States share many life history traits and demonstrate complex interspecific interactions that range geographically from competitive exclusion to sympatry. P. jordani was introduced to Mountain Lake Biological Station, Virginia, USA, between the years 1935 and 1945. We tested whether competition for space may influence the invasion of P. jordani into native P. glutinosus habitat by utilizing data from natural distributions, a field experiment, and controlled laboratory experiments. No environmental variables differed where P. glutinosus and P. jordani were collected in the field at the site of P. jordani introduction. In the field experiment, P. glutinosus was more fully exposed during foraging bouts in cages shared with heterospecifics as opposed to ones shared with conspecific salamanders. Condition (mass relative to body length) of salamanders at the end of the 3 months did not differ between conspecific and heterospecific treatments. In the laboratory, P. glutinosus most often attained the single burrow in the arena, but residency status had no effect. Species cohabited the burrow 50% of the time. Pair-wise encounters in the laboratory indicated that both species spend less than 20% of the time in aggressive behaviors as juveniles. Adults showed no behavior interpreted by us as aggression during pair-wise encounters. Received: 19 December 1999 / Accepted: 18 March 2000     

Nitrate- and silicate-competition among antarctic phytoplankton

Natural phytoplankton from antarctic waters in the Drake Passage were used for competition experiments in semicontinuous cultures. The outcome of interspecific competition for silicate and nitrate was studied at a range of Si:N ratios (from 2.6:1 to 425:1) and at three different dilution rates. For five species Monod kinetics of silicate-and nitrate-limited growth has been established. Comparison between theoretical predictions derived from Monod kinetics and the outcome of competition experiments showed only minor deviations. Contrary to literature data, considerable depletion of nitrate was found in antarctic seawater. Both the concentrations of soluble silicate and of nitrate were too low to support maximum growth rates of some of the diatom species under investigation.     

Sympatric sexual signal divergence among North American Calopteryx damselflies is correlated with increased intra- and interspecific male–male aggression

Divergence of sexual signals in sympatry can arise as a consequence of (1) interspecific competition for resources, (2) selection against maladaptive hybridization, or (3) as a result of selection to reduce the cost of interspecific aggression; termed agonistic character displacement (ACD). Calopterygid damselflies have emerged as a model system for studying the evolution of divergent sexual signals due to the repeated evolution of sympatric species pairs with fully and partially melanized wings. Damselfly wing patterns function during both courtship and territory defense. However, the relative contributions of natural and sexual selection to phenotypic divergence and enhanced isolation in sympatry remain unclear in many cases. Here, we investigated the hypothesis that interference competition, in the form of increased interspecific male–male aggression, drives the evolution of character displacement in sympatry between two species of North American damselflies, Calopteryx aequabilis and Calopteryx maculata, that show no evidence of ecological divergence or ongoing hybridization. In paired behavioral trials, we found that interspecific male aggression related to territory defense varied between site, species, and as a function of the relative abundance of con- vs. hetero-specific males. Specifically, we found that large-spotted C. aequabilis males received increased intra- and interspecific aggression but that aggression against large-spotted males declined during the middle of the flight season when both species were equally abundant. Based on these results, we suggest that ACD leads to enhanced species recognition, and may be a common outcome of the antagonism between interspecific male–male competition and the countervailing force of intraspecific sexual selection favoring increased wing melanization among territorial damselfly species.