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.     

No male agonistic experience effect on pre-copulatory mate choice in female earwigs

Mating with dominant males may confer considerable benefits, but also incur significant costs, hence intrasexual competitiveness is a likely target of mate choice. In addition to established modes of mate assessment, females may use cues or signals associated with agonistic experience effects to assess the relative competiveness of males. Experience effects, where the outcome of a fight increases the likelihood of a similar outcome in subsequent fights, may result from an animal’s altered state after conflict, but can also arise from strategic rival use of information perceived about this altered state. While females may similarly use this information in mate choice decisions, this potential consequence of male–male conflict has largely been neglected. Here, we investigate the effects of experience on subsequent agonistic performance in the earwig Euborellia brunneri by imposing winning or losing experiences on males and rematching them with naïve, size-matched rivals. We reveal a strong loser effect in subsequent fights, with nearly all previous “losers” losing against new rivals. In contrast, we found no equivalent winner effect, with previous “winners” exhibiting no increased likelihood of winning. We subsequently test whether the effects of male agonistic experience extend to pre-copulatory female mate choice. We show that females, when allowed to choose between naïve males and “winners” or “losers”, do not discriminate between males based on their recent agonistic experience. Therefore, while fighting history can play an important role in male–male interactions, females may not attend to this information.     

The role of size and aggression in intrasexual male competition in a social lizard species, Egernia whitii

Competition between males is a key component of the agonistic intrasexual interactions that influence resource acquisition, social system dynamics, and ultimately reproductive success. Sexual selection theory predicts that traits that enhance success in intrasexual competition (particularly male–male competition) should be favored. In vertebrates, this often includes body size and aggression, with larger and/or more aggressive males outcompeting smaller or less aggressive conspecifics. The majority of studies consider aggression as a flexible trait which responds to local social or environmental conditions. However, aggression frequently shows considerable within-individual consistency (i.e., individuals have identifiable aggressive behavioral types). Little is known about how such consistency in aggression may influence competition outcomes. We integrated a detailed field study with a laboratory experiment to examine how a male’s aggressive phenotype and his size influence competitive interactions in Egernia whitii, a social lizard species which exhibits strong competition over resources (limited permanent shelter sites and basking sites). Individual aggression and size did not predict competition outcome in the laboratory nor did they predict home range size, overlap, or reproductive success in the field. However, winners of laboratory trial contests maintained consistent aggressive phenotypes while consistency in aggression was lost in losers. We suggest that aggression may be important in other functional contexts, such as parental care, and that alternative traits, such as fighting experience, may be important in determining competition outcome in this species.     

Signaling effort does not predict aggressiveness in male spring field crickets

The relationship between signaling and aggression is still unclear despite several decades of research. However, there is a growing interest in studying signals that predict aggressive behavior or fighting ability. The goals of our study were threefold: we investigated the relationship between signaling effort and aggression, the relationship between body condition and aggression, and the effect of fighting experience on subsequent signaling behavior in wild-caught and laboratory-reared male spring field crickets (Gryllus veletis). We found that aggressive behavior was not related to signaling effort, body size, or body condition. For contest winners, wild-captured males were more aggressive than laboratory-reared males. Signaling effort was highly repeatable within individuals, but aggressive behavior had low repeatability. We found no evidence for a winner or loser effect on signaling; there was no change in signaling effort when we compared contest winners and losers before and after they participated in aggressive contests. Long-distance acoustic signaling and aggressive behavior appear to be independent of one another in spring field crickets, perhaps serving different functions in female attraction and male–male competition, respectively.     

Male Iberian rock lizards may reduce the costs of fighting by scent matching of the resource holders

Animals should adopt strategies to minimize the costs of intraspecific aggressive interactions. For example, individuals should be able to identify resource holders in advance and avoid fighting with them because residents are generally more likely than intruders escalate aggression. It has been suggested that scent marks function mainly to allow competitor assessment by conveying the costs of entering a scent-marked area. Individuals may identify territory owners by comparing the scent of substrate marks with the scent of any conspecific they encounter nearby, assessing whether these two scents match or not, a mechanism known as scent matching. Here, we examined the response of male Iberolacerta cyreni lizards to areas scent-marked by other males and the potential role of scent matching in agonistic interactions. We designed a laboratory experiment where we allowed a male to explore the scent-marked substrate of another male, and then we immediately staged agonistic encounters in a nearby clean neutral area with either the male that had produced the scent marks (matching treatment) or with a different non-matching individual male. The higher chemosensory exploratory rates of substrate scent marks in comparison to clean substrates suggested that males detected and spent more time exploring scent marks to obtain information on the donor male. Moreover, this information was later used to decide the fighting strategy. Intruding males delayed time until the first agonistic interaction, reduced the intensity of fights and the number of aggressive interactions, and won less interactions with males which scent matched that of scent marks (because they would be considered as the territory owners) than with other non-matching individuals. Our results show that male I. cyreni lizards use scent matching as a mechanism to assess the ownership status of other males, which could contribute to modulate intrasexual aggression, reducing costs of agonistic interactions.     

Desperado siblings: uncontrollably aggressive junior chicks

According to the desperado sibling hypothesis, chicks of obligately siblicidal species kill their junior broodmates as early as possible because junior broodmates face dire ecological prospects and are consequently predisposed to aggressively overthrow elder broodmates by all-out aggression. The agonistic behavior of junior chicks is virtually unknown because they die so young. We investigated the agonistic behavior of hypothetical desperados by fostering junior brown boobies (Sula leucogaster, an obligately siblicidal species) into nests of the blue-footed booby (S. nebouxii, a facultatively siblicidal species) containing an older singleton blue-foot chick. Controls were junior blue-footed boobies fostered into the same situation. Junior brown boobies were 7 times more aggressive than controls and most of them dominated their elder and larger nestmates. Four of nine brown booby juniors showed relentless aggression, delivering up to 711 pecks, bites and pushes (including "expulsion pushes") per hour, thereby overwhelming nestmates 90% heavier and permanently expelling one of them from the nest. Similarly, in natural broods of two surviving brown booby chicks, the losing chick was 13 times as aggressive as blue-foot subordinates, up to at least age 7 weeks. Contrast of the two species of booby suggests the evolution of agonistic roles within broods may be partly driven by selection on potential victims to express a level and type of aggressiveness appropriate to their status-related ecological prospects.     

Reproductive competition and fecal testosterone in wild male giant pandas (<Emphasis Type="Italic">Ailuropoda melanoleuca</Emphasis>)

The giant panda (Ailuropoda melanoleuca) is notoriously difficult to study in the wild, but its interesting reproductive ecology makes the effort worthwhile. Perhaps more than most species, the panda is energy-limited, which alters the cost/benefit analysis of its reproductive ecology. Using global positioning system/very high frequency radiocollars to locate mating aggregations, we used behavioral observations and fecal testosterone assays to gain insight into male panda reproductive effort and strategies, and test theories relating to reproductive competition. Male pandas initially competed fiercely for access to females that were about to be fertile, but once male competitive status was determined, aggression rates declined. Contact aggression was only observed during the first 2 days of mating aggregations; thereafter, it was replaced with noncontact aggression and avoidance. Agonistic interactions were highly asymmetrical, with contest losers (subordinates) showing less aggression and more avoidance than contest winners (dominants), both before and after contest outcome was established. The competitively superior male displayed mate-guarding tactics and secured all observed copulations. Contrary to theoretical predictions, testosterone levels did not predict aggression levels or contest winners and also were not affected by winning or losing a contest. Body size appeared to be the primary determinant of contest outcome. We discuss our findings in light of theoretical predictions, such as those arising from the “challenge hypothesis,” in the context of the giant panda’s foraging and nutritional ecology.     

The formation and maintenance of crayfish hierarchies: behavioral and self-structuring properties

Behavioral changes in fighting and the development of dominance relationships were analyzed in groups of juvenile crayfish (Astacus astacus) using quantitative behavioral techniques. When individuals were placed into an aquarium, the number of agonistic challenges, their mean duration, and maximum intensity reached were high initially but then decreased steadily as the hierarchy developed. In all groups, linear hierarchies emerged which became increasingly stable over time. Winning influenced subsequent fighting behavior on two distinct time scales. In the short term, recent winners became progressively less likely to retreat. Second, individuals occupying dominant positions for days became increasingly likely to escalate to higher intensities early in the encounter. Both effects biased the outcome of future interactions such that winning enhanced further success and losing decreased an individual’s subsequent chances for dominance. Received: 11 November 1999 / Revised: 1 March 2000 / Accepted: 3 May 2000     

Delayed plumage maturation in the orchard oriole (Icterus spurius): tests of winter adaptation hypotheses

Summary Explanations of delayed plumage maturation (DPM) in passerines have focused on potential breeding season advantages for non-definitive (subadult) plumage. In contrast, the molt constraint hypothesis (Rohwer and Butcher 1988) proposes that subadult plumage is a winter adaptation, increasing winter survivorship by decreasing intraspecific aggression (the winter status signaling hypothesis) or predation (the winter crypsis hypothesis). Under the molt constraint hypothesis, nondefinitive breeding plumage is non-adaptive, resulting from species-specific constraints on the number of feathers replaced during the pre-alternate (spring) molt. In studies conducted on an overwintering population of orchard orioles in Panama, I tested predictions of both the winter status signaling and molt constraint hypotheses. Contrary to predictions of winter status signaling, I found no evidence that subadult plumage reduces adult male aggression toward subadults. Agonistic encounters occurred at random with respect to plumage and the intensity of adult male/subadult male encounters was not lower than the intensity of encounters occurring within age classes. Contrary to the molt constraint hypothesis, I found no evidence that the number of feathers exchanged by subadults in their pre-alternate molt is the sole constraint on the development of subadult breeding plumage. The majority of feathers grown by molting subadult orioles during January and February were of non-definitive coloration. These results, together with results of earlier breeding season experiments which tested summer communication hypotheses for DPM in this species, suggest that subadult plumage in the orchard oriole may be non-adaptive and the result of constraints on plumage development. They also indicate, however, that the extent of the pre-alternate molt is not the sole source of that constraint.     

Causes and consequences of contest outcome: aggressiveness, dominance and growth in the sheepshead swordtail, Xiphophorus birchmanni

Although our understanding of how animal personality affects fitness is incomplete, one general hypothesis is that personality traits (e.g. boldness and aggressiveness) contribute to competitive ability. If so, then under resource limitation, personality differences will generate variation in life history traits crucial to fitness, like growth. Here, we test this idea using data from same-sex dyadic interaction trials of sheepshead swordtails (Xiphophorus birchmanni). In males, there was evidence of repeatable variation across a suite of agonistic contest behaviours, while repeatable opponent effects on focal behaviour were also detected. A single vector explains 80 % of the among-individual variance in multivariate phenotype and can be viewed as aggressiveness. We also find that aggressiveness predicts dominance—the repeatable tendency to win food in competition—and dominant individuals show faster post-trial weight gain (independently of initial size). In females, a dominance hierarchy predictive of weight gain was also found, but there was no evidence of variation in aggressiveness. While size often predicts contest outcome, our results show that individuals may sometimes grow larger because they are behaviourally dominant rather than vice versa. When resources are limited, personality traits such as aggression can influence growth, life history, and fitness through impacts on resource acquisition.     

When to be neighbourly: differential agonistic responses in the lizard Platysaurus broadleyi

Differential treatment of neighbours and strangers (the dear enemy phenomenon) can reduce the costly effects of territory defence. Individual recognition, and by extension the dear enemy phenomenon, was examined in the cordylid lizard Platysaurus broadleyi. I had no knowledge of familiarity between individuals and therefore classed intruders as neighbours or non-neighbours and tested for differences in agonistic response consistent with the dear enemy phenomenon. In the dear enemy phenomenon, levels of aggression are inversely related to degree of familiarity such that residents are more tolerant of immediate neighbours than distant territory holders. A manipulative field experiment in conjunction with field observations of known territory holders revealed that resident males allowed neighbours to approach more closely than non-neighbours before challenging them, residents were more aggressive towards non-neighbours than neighbours, contests between neighbours were significantly longer than between neighbours and non-neighbours and contests between residents and non-neighbours were likely to result in a win for the resident, while neighbours frequently drew contests. These results suggest a level of recognition consistent with the dear enemy phenomenon. I also manipulated front leg colour to test the hypothesis that it alone could serve as a cue for individual recognition. I found no support for this hypothesis and suggest that multiple cues operate in individual recognition. Received: 4 March 1999 / Received in revised form: 11 April 1999 / Accepted: 19 April 1999     

Why do winners keep winning?

Summary Winners of aggressive interactions often continue to win in future encounters. I propose that this phenomenon is a by-product of two other phenomena occurring simultaneously: first, initiators of aggressive interactions typically win those interactions and, second, winners of aggressive encounters often become more likely to initiate future interactions. I propose that these phenomena can be explained, in turn, by selection favoring an individual's initiating an interaction only when it is likely to win that interaction. I found support for this two-step explanation for the winning begets winning phenomenon by observing aggressive interactions among captive dark-eyed juncos (Junco hyemalis oreganus). First, initiators of interactions almost always won. Second, based on characteristics of winners in birds' home aviaries, I could predict which birds would initiate against novel competitors — winners were long-winged males with dark hoods, and birds with these characteristics were more likely to be the first to initiate in novel triads. In addition, aggression was directed preferentially towards other dark-hooded males. The results of this study may expand our understanding of the dynamics of aggressive interactions.     

Intraspecific aggression in the colour morphs of the anemone Phymactis clematis from Chile

The anemone Phymactis clematis exhibits intraspecific agression. An investigation into dominance relationships amongst the 3 colour morphs collected from Vna del Mar, Chile, during June, 1980, revealed them to be equally aggressive. The outcomes of experimental contests appeared to reside solely with size-dependent differences in the threshold for the release of agression. Contests were asymmetric and larger individuals displayed aggression earlier against smaller opponents, and won encounters. An analysis of times for the onset of aggression in contests showed that they were highly variable and were not a good indicator of the absolute aggressive ability of individuals as denoted by dominance rankings. It provided, however, evidence indicating a lack of assessment of opponent size during initial contact. This is discussed in relation to the evolution of aggression in anemones and to the concept of evolutionarily stable strategies in contest behaviour. It is concluded that future work on the apparent size-dependent threshold will most instructively be conducted at the neurophysiological level.     

The timing of chemical signaling with urine in dominance fights of male lobsters (Homarus americanus)

Chemicals carried in the urine are a rich source of information about the identity, sex, aggressive motivation, and other attributes of an animal. In agonistic interactions, animals should be careful about disclosing such information, since it can be used by receivers to the disadvantage of the sender. By adjusting the timing of urine release, the signaler may still influence the behavior of the receiver to its own benefit. Here we investigate the urine signaling of American lobsters (Homarus americanus), necessary for the maintenance of dominance, using a catheter technique with high temporal resolution. We hypothesize that urine release in lobster fights is not continuous but restricted to moments that may elicit a reaction in the receiver beneficial to the sender. We found that eventual winners signaled more readily in the initial phase of the fight. Eventual losers did not show such initial urination. This suggests that a confident lobster may use urine signals to influence the behavior of its opponent into giving up the fight at an early stage. For both winners and losers, urine release is linked to offensive behavior and increases as the intensity of agonistic behavior increases. The coupling of urine release to offensive behaviors appears to improve both the reliability and efficacy of the chemical signal. Releasing urine under the increasing risk of inflicting injury during the fight provides a handicap that may ensure reliability of the threat signal. Furthermore, the immediate coupling of urine components to offensive behavior may help to consolidate the receiver"s memory of the signaler"s individual scent and thus facilitate future recognition of the dominant animal. Received: 6 October 1999 / Accepted 28 August 2000     

Do Midas cichlids win through prowess or daring? It depends

Summary Individual Midas cichlids (Cichlasoma citrinellum) show persistent differences in aggressive behavior toward dummy fish. The starting question was whether the level of such behavior can be used to predict the winner of a fight. In the first (long-term) experiment two fish were matched for size, color, and sex. After 24 h the opaque barrier separating the 2 fish was removed; they immediately behaved aggressively. Scores for aggression toward dummies did not predict the winners, nor did taking the initiative in escalation. But weight did foretell the winners, who averaged only 2% heavier than their opponents. The conventional display phase of such fights was brief (20% of total duration), and escalation was rapid. Both winners and losers sustained damage, but losers accumulated damage faster than winners. In the subsequent (short-term) experiment the fish were separated only 1–2 h. Now aggression scores predicted winners, and winners were the fish who escalated. Weight of fish had no effect. The conventional phase was relatively much longer, about half the length of the fight. Losers accumulated damage at the same rate as the losers in the long-term experiment, but the fights were shorter; winners suffered little damage. The fish had difficulty assessing one another. Fighting prowess was remarkably uniform when weight was factored out. Daring to escalate, in contrast, varied among individuals and correlated with aggression scores. Prowess (=weight) determined the outcome in the long-term experiment, which may find its parallel in nature in intraterritorial disputes. Daring to escalate determined the winner in the short-term experiment; this may be comparable to establishing a territory in nature.Prowess probably results from strong directional selection because it has low costs and high benefits. In contrast, daring is subject to bi-directional selection because both costs and benefits are high. Resource holding potential is conventionally viewed as deriving from prowess of self and opponent and value of resource; to that one must add the individual's inherent aggressiveness.     

The appeasement effect of sterility signaling in dominance contests among Bombus terrestris workers

The establishment of dominance hierarchies through aggressive interactions is very common in insect societies. In many cases, it is also mediated through pheromone emissions that enable individuals to evaluate the reproductive quality and level of aggressiveness of the dominant individual, thereby reducing the number and intensity of costly fights. Here, we studied these processes in the primitively eusocial bee Bombus terrestris, using a paired bee system. Specifically, we investigated the behavioral, reproductive, and pheromonal correlates of dominance establishment. Workers were shown to establish dominance hierarchies using overt aggression within 3–4 days. Thereafter, the aggression drastically decreased, and dominance was maintained mostly by ritualized agonistic behavior. The behaviorally dominant bee lost the ester compounds that workers produce in their Dufour's gland (the so-called “sterility signal”) concomitantly with the development of her ovaries. The other bee announced as subordinate by continuously producing high amounts of those esters. The hypothesis that sterility signaling serves as an appeasement signal to pacify the dominant bees is supported by the negative correlation found between the proportion of these esters and the level of aggression that the subordinate received from the dominant worker. Physical interactions, and presumably also the ensuing overt aggression between the bees, were essential for the above pheromonal change to take place and enabled the dominant workers to develop their ovaries and to lay eggs. The subordinate bee’s signaling of non-reproductive status may minimize energy expenditure in costly fights and help stabilize the reproductive division of labor among workers.     

Regional variation in queen and worker aggression in incipient colonies of the desert ant Messor pergandei

The fate of queen foundress associations in ants varies across taxa: in some, lethal fighting results in survival of a single queen, while in others, queens coexist long term. One hypothesis for this difference is that selection favors fighting when group sizes are small and tolerance when groups are large. In an experiment with the ant Messor pergandei, we formed small, medium, and large groups with newly mated queens from field populations that have different mean group sizes and differ in whether multiple queens occur in older established colonies. We found that whether queens are eliminated by fighting depends upon region of origin and not group size: regardless of co-foundress number, queens from sites with single-queen adult field colonies displayed agonistic behaviors and their colonies reduced to a single queen, while queens from sites with multiple-queen colonies did not fight and co-foundresses coexisted long term. Worker aggression towards and elimination of queens were also correlated with region of origin. Where fighting occurred, queens were as likely to be killed by workers as by other queens. An aggressive display was the most common form of agonistic interaction among queens, while fighting was relatively rare. We hypothesize that queen displays evolved in response to worker attacks because they increase the probability that workers will eliminate competitor queens. Our results suggest that the evolutionary interests of workers, as well as queens, could be important in determining the evolution and maintenance of queen elimination in foundress associations.     

Information content is more important than sensory system or physical distance in guiding the long-term evolutionary relationships between signaling modalities in Sceloporus lizards

Long-term signal evolution is shaped by a variety of selective pressures including the need to convey additional information or to improve message transfer to specific receivers or through multiple environments. Here, we test the relative importance of information and sensory modality in shaping the long-term evolution of multimodal signals in Sceloporus lizards. To broadcast identity at territorial boundaries, male Sceloporus use both visual motion (headbob) and chemical signals, whereas they use color (blue belly patches) to signal aggression. Using modern phylogenetic comparative methods, we found a negative correlation between evolutionary changes in visual motion (headbobs) and chemical (femoral pore) signals, but only indirect ties between the evolution of color and motion signals (both of which are perceived visually) through viviparity, and no evidence of an evolutionary link between color and chemical signals. We also find a negative correlation between arboreality and chemical signals. Thus, information content (in this case, broadcasting individual identity versus signaling aggression) appears to play a more important role than sensory modality or physical distance in guiding long-term signal evolution. Additional insights into the underlying evolutionary processes are described, illustrating the utility of a phylogenetic approach.     

Community composition and nearest neighbor relationships in orb-weaving spiders: the product of aggression?

Summary Nearest-neighbor distances and species' densities were calculated for 949 individuals of 8 species of orb-weaving spiders from eight edge and meacow sites in Panama. These data were used to test the hypothesis that interspecific aggressive interactions influenced spider community structure by: (1) eliminating or decreasing the densities of sympatric, less aggressive species, or, (2) affecting nearest-neighbor relationships so that large or aggressive individuals had smaller, less aggressive or more distant neighbors. Although there was a negative relationship between the densities of two species pairs (Leucauge sp. and Mangora pia; Eriophora fuliginea and Cyclosa caroli), overall spider densities could not be predicted from indices of spider aggression obtained by introduction experiments. The spatial distribution of spiders in three dimensions was not significantly different than random. I attributed the lack of clumping to the relative vegetational homogeneity of my sites; the lack of overdispersion may reflect the fact that websites are not limiting. Relationships between nearest neighbors generally supported the aggression hypothesis, however, comparisons of the actual data with equivalent randomly-assorted data indicate that a hypothesis of randomly chosen neighborsis equally applicable. This result suggests that, unless websites differ significantly in their quality, with high quality websites in short supply, spider aggression may not affect community structure.     

Begging behavior and food acquisition by brown-headed cowbird nestlings

Understanding the selective forces that limit the exaggeration of begging signals is a critical issue in understanding the evolution of begging behavior. I studied the begging behavior of nestlings of the brown-headed cowbird (Molothrus ater), a brood parasite. In the nests of indigo buntings (Passerina cyanea), brown-headed cowbird nestlings received approximately twice as much food per hour than their host nestmates. I tested three hypotheses for the mechanism by which cowbirds acquired more food than their bunting nestmates: the size advantage hypothesis, the signal exaggeration hypothesis, and the novel begging behavior hypothesis. I found support for the hypotheses that cowbirds acquire more food as a result of their larger body size, and due to the exaggeration of begging signals that are not dependent on body size. I did not find support for the role of novel begging behaviors in cowbird food acquisition. These results suggest that food acquisition by host chicks in unparasitized nests could be increased by the exaggeration of begging signals. Recent work suggests that such exaggeration may be limited by the risk of nest predation, but further studies are needed. Received: 12 December 1997 / Accepted after revision: 29 December 1997