Fighting rules and rival recognition reduce costs of aggression in male lizards, Podarcis hispanica

The establishment of fighting rules and the ability to recognise individual conspecifics and to assess their fighting ability and/or roles may help to reduce costs of fighting. We staged encounters between males of the lizard Podarcis hispanica to examine whether lizards used fighting strategies and whether a previous agonistic experience affects the outcome and characteristics of a subsequent encounter. The results showed that simple rules such as body size differences and residence condition were used to determine the outcome of agonistic interactions as quickly as possible. Thus, larger males were dominant in most encounters. However, when size differences between opponents are smaller, they may be more difficult to estimate and, then, residence condition was more important. In addition, the intensity of interactions between males could be explained according to the ”sequential assessment game”, supporting the idea that P. hispanica males acquire information about fighting ability gradually during the progress of a fight. Our results also showed that the second fight of the same pair of males was less aggressive, even when its outcome was the opposite of the first. This result suggests that male P. hispanica can recognise individual opponents and that they use this information to reach a contest outcome more quickly, thus reducing unnecessary aggression levels in subsequent interactions. These fighting strategies and assessment mechanisms may help to stabilise the social system of this lizard. Received: 2 November 1999 / Revised: 26 August 2000 / Accepted: 4 September 2000     

Sexual selection and mating system in Zorotypus gurneyi Choe (Insecta: Zoraptera)

Body size is clearly an important factor influencing the outcome of agonistic contests, but is often weakly correlated with dominance ranks in Zorotypus gurneyi Choe (Insecta: Zoraptera). The study of the development and dynamics of dominance relations using artificially constructed colonies show that age, or tenure within the colony, is the prime determinant of dominance among males. Dominance hierarchies become relatively stable within 2 or 3 days and males that emerge later normally begin at the bottom of the hierarchy regardless of size. Males interact much more frequently when they are simultaneously introduced to each other than when they are allowed to emerge at different times. In the latter case, males that emerge late appear to recognize relative dominance of older males and avoid direct contests. Considering the high correlation between dominance rank and mating success, there is a strong selective advantage to males that emerge earlier and such pressure of sexual selection may be responsible for the difference in life history strategies between Z. gurneyi and its sympatric congener, Z. barberi Gurney, in central Panama.     

Asymmetrical contests in defence of rock ptarmigan territories

Summary Contests involving territorial male rock ptarmigan (Lagopus mutus) occur frequently over the course of the breeding season. Fights are initiated when one male (Neighbour or Bachelor) intrudes on another's (Resident) territory. These contests involved two kinds of asymmetries: Neighbours were similar to Residents in mating status, size and experience, differing only in the gains accrued from winning a fight (payoff asymmetry) whereas Bachelors were smaller, less experienced males that had no mate or territory (fighting asymmetry, in addition to a relatively larger payoff asymmetry than in Resident x Neighbour contests). In this study, we describe and compare the contests resulting from these intrusions, and we assess how the asymmetries (fighting ability, payoffs) between opponents influenced contest intensity. Resident x Neighbour contests were brief, highly-ritualized affairs, while Resident x Bachelor interactions were long, intense battles, often ending in direct physical attack. These findings are in direct opposition to many theoretical and empirical studies on fighting asymmetries which suggest that contest intensity should be negatively correlated with the degree of fighting asymmetry between contestants. Since the potential losses associated with a contest differed between contestants and were highest for an intruding Neighbour, we conclude that payoff asymmetries were more important than fighting asymmetries in influencing a male's tendency to escalate.     

Chemical rival recognition decreases aggression levels in male Iberian wall lizards, <Emphasis Type="Italic">Podarcis hispanica</Emphasis>

In an agonistic interaction, the assessment of the probable outcome of future encounters with the same individuals may be the best way of decreasing costs of fighting, but this may only be accomplished if animals are able to recognize individual conspecifics. We staged encounters between male lizards, Podarcis hispanica, to examine whether odoriferous cues are involved in rival recognition during agonistic interactions. We experimentally manipulated the odour of intruding males, creating familiar males with their own odour or bearing odours of unfamiliar males, and unfamiliar males bearing unfamiliar odours or odours of familiar males. The results showed that when familiar males were impregnated with scents from unfamiliar males, they elicited an aggressive response by resident unmanipulated males similar to that observed for a new unfamiliar male with unfamiliar odour. This suggests that resident males were unable to recognize familiar males when their own scents were removed. In contrast, responding males were less aggressive towards familiar males impregnated with their own odour and towards unfamiliar males impregnated with scents of familiar males, suggesting that when two males have already interacted, their scents become familiar for both males, and that the detection in successive encounters of the familiar scent suffices to reduce the aggressive response of territorial males. Therefore, recognition mechanisms based on chemical cues during agonistic encounters may contribute to reducing the intensity and the costs of fighting in P. hispanica and may play an important role in the organization of their social system.     

Roles of male residence and relative size in the social behavior of Iberian rock lizards, Lacerta monticola

Many studies on contest competition used residency asymmetry as a discrete variable. However, the probability of winning an interaction may change as a continuous function of the value of the location where the encounter occurs. We performed a field study to examine the importance of location within a home range and relative body size to the outcomes of agonistic interactions between male lizards, Lacerta monticola. The distances to activity centers (the most used locations based on a density function of sightings) and relative size play important roles in agonistic interactions and had interacting effects in natural conditions. On the other hand, previous studies with lizards suggested that inferior competitors are able to avoid agonistic interactions in the field. Thus, we staged encounters in the laboratory to examine the behavioral responses of smaller individuals. The responses of each focal smaller male were measured in its own home cage (resident), in the cage of a larger male (intruder) and in a cage in which no male was previously present (control). The predominant behavioral tactics of smaller males were avoidance when they are the intruders and displaying when they are the residents. Submissive displays by smaller males may help reduce the costs of agonistic encounters.     

Chelipeds are the real weapon: cheliped size is a more effective determinant than body size in male–male competition for mates in a hermit crab

To understand the evolution of weapons, we must understand both their functions and relative importance compared to body size in determining fighting success. Many decapod crustaceans develop disproportionately large chelipeds for their body size and use them as a weapon in agonistic interaction. There are, however, examples where weapons are merely signals of resource holding potential (RHP) and the RHP is actually determined by body size. We investigated the function and relative efficacy of body size and major cheliped size in male–male contests for females in the hermit crab Diogenes nitidimanus. Contests over females took two forms: (1) males preemptively guarded females and opponents did not fight with the guarding male. Cheliped size contributed significantly to the settlement of these contests and probably functioned as a visual signal for the opponents. (2) Guarding males engaged in physical combat with an opponent. In these cases, both body and cheliped sizes affected contest outcomes. The effect size for cheliped size was as strong, or stronger, than that for body size. These results suggest that large chelipeds have evolved as a true weapon and are effective in escalated fights for resources. Therefore they are also efficient visual signals for settling contests with only display. Our results are a rare example that clearly demonstrate that weapons are a more important determinant of fights than body size when both body and weapon size affect resource acquisition.     

“Alternative” mating behavior in the swordtails Xiphophorus nigrensis and Xiphophorus pygmaeus (Pisces: Poeciliidae)

Summary The swordtail Xiphophorus nigrensis exhibits three relatively discrete male body-size classes that derive from allelic variation at the Y-linked pituitary (P) locus. Previous studies have shown that larger males have greater relative reproductive success, and that females prefer large males. We describe the mating behavior utilized by males of each size class during individual encounters with females. Small males rely on chase behavior, similar to the alternative mating behavior classified as sneaker in small males of other species. Large males court and intermediate-sized males court or chase, depending on their body size. There is a strong correlation between P alleles for small size (s) and large size (L) with chasing and courting, respectively. The relationship between mating behavior of males of the genotype I is ambiguous. In the closely related species X. pygmaeus, males are of size similar to smaller X. nigrensis males. Paradoxically, these males do not show the courtship display that typifies larger male X. nigrensis and many other species of swordtails, but instead often employ chase behavior identical to the alternative mating behavior in small male X. nigrensis. We suggest that historical and genetic constraints, in addition to current selection forces, might be important factors in explaining the existence of alternative mating behavior in X. pygmaeus.     

Exaggerated male eye span influences contest outcome in stalk-eyed flies (Diopsidae)

Evolution of male weapons or status signals has been hypothesized to precede evolution of female mating preferences for those traits. We used staged male fights among three species of Malaysian stalk-eyed flies (Diptera: Diopsidae) to determine if elongated eye span, which is preferred by females in two sexually dimorphic species, influences contest outcome. Extreme sexual dimorphism, with large males possessing longer eye span than females, is shared by Cyrtodiopsis whitei and C. dalmanni. In contrast, C. quinqueguttata exhibits a more ancestral condition – short, sexually monomorphic eye stalks. Videotape analysis of 20-min paired contests revealed that males with larger eye span and body size won more fights in the dimorphic, but not monomorphic, species. To determine if males from the dimorphic species use eye span directly to resolve contests, we competed male C. dalmanni from lines that had undergone artificial selection for 30 generations to increase or decrease eye span. We found that eye span, independently of body size, determines contest outcome in selected-line males. Furthermore, in both dimorphic species, the average encounter duration declined as the eye span difference between contestants increased, as expected if males use eye span to assess opponent size. The number of encounters also increased with age in dimorphic, but not monomorphic, species. Selected-line males did not differ from outbred males in either fight duration or number of encounters. We conclude that exaggerated male eye stalks evolved to influence both competitive interactions and female mating preferences in these spectacular flies. Received: 20 July 1998 / Received in revised form: 2 February 1999 / Accepted: 12 March 1999     

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.     

Dominant male red junglefowl (Gallus gallus) test the dominance status of other males

In sexual selection, honest signals are maintained by a variety of mechanisms. In red junglefowl (Gallus gallus), health, condition and social status affect comb size, a well-documented predictor of female choice. The comb size of subordinate male junglefowl appears to be suppressed when in the company of other males. One hypothesis for how social status could affect ornament expression in this way involves punishment of cheaters. Under this scenario, dominant males periodically challenge similar males signalling putative high status. For subordinate males, the risk of fighting a high-ranked male could make it prohibitively costly to develop ornamentation signalling dominance. We asked if dominance signals influenced the direction of aggression by dominant males. To address this issue, we conducted experiments in which 19 dominant-acting, large-combed male junglefowl were allowed to choose to fight one of two opponents. The two potential fight opponents differed in comb size, dominance behaviour, or in both traits. In 15 of 19 trials, dominant-acting males chose to fight large-combed, dominant-acting opponents rather than small-combed, subordinate-acting opponents. This is the first demonstration that aggression of dominant male birds is directed at other males based on the display of an ornament known to be attractive to females. However, males did not discriminate between fight opponents when potential opponents differed in only one of the two status indicators (large-combed males chosen in 11 of 19 trials, dominant-acting males chosen in 10 of 19 trials).     

Weapon size versus body size as a predictor of winning in fights between shore crabs, Carcinus maenas (L.)

Relative body size (carapace width) and weapon size (chela length) were used as indicators of resource holding potential (RHP) in the agonistic behaviour of male shore crabs, Carcinus maenas (L.). Weapon size was found to be a more reliable predictor of the outcome of pairwise fights than body size. Crabs with longer chelae than their opponents were more likely to win fights than crabs with relatively larger bodies. Body size had less influence on the outcome of fights. Relative body and weapon size did not influence initiation of contests but did affect the likelihood of winning; however, this was significant only for weapon size. Winning crabs had heavier claws with greater surface area than losing crabs. There was no relationship between relative size and fight duration. The frequency of cheliped display increased with chela length and win- ners performed significantly more displays than losers. Received: 5 February 1997 / Accepted after revision: 20 May 1997     

Differential attraction of females to dominant males: Olfactory discrimination and mating preference in the brown lemming (Lemmus trimucronatus)

Summary This study investigated differential attraction of estrous brown lemmings (Lemmus trimucronatus) to conspecific males recently exposed to each other for a 10-min agonistic encounter. In tests conducted 5 min, 1 h and 24 h after agonistic encounters, females preferred the oder of dominant males to that of defeated males when both odors were presented simultaneously in a Y-maze olfactometer. Defeat in an agonistic encounter did not reduce the propensity of male lemmings to initiate sexual behavior. In one-male, one-female tests conducted 5 min after agonistic encounters, dominant males achieved higher mount and thrust scores while defeated males obtained higher scores for attempted mounts. The sexual behavior of dominant and defeated males did not differ significantly in similar tests conducted 1 h and 24 h later. In contrast, females readily mated with dominant males and tended to avoid defeated males in two-male tethering tests conducted 5 min after agonistic encounters. In these tests, females still showed a preference for dominant males 1 h and 24 h after male agonistic encounters.     

The role of body size and fighting experience in predicting contest behaviour in the black field cricket, <Emphasis Type="Italic">Teleogryllus commodus</Emphasis>

Body size strongly predicts fighting behaviour and outcome in many species, with the larger opponent usually winning contests. However, recent fighting experience can have a strong influence on the establishment of dominance hierarchies, with recent winners being more likely to win subsequent contests, while recent losers are more likely to lose. Recent fighting experience therefore has the potential to modify the effect of body size in determining contest behaviour and outcome. Here, we investigate whether recent fighting experience weakens the role of body size in predicting contest behaviour in the black field cricket, Teleogryllus commodus. We compared the role of body size in determining contest outcome during initial non-physical encounters and escalated aggressive physical encounters (grapples), as well as the probability of escalation occurring, in contests involving either a naïve or experienced smaller male against a naïve larger male. We found that recent fighting experience only affected contest outcome during non-physical encounters. Once a contest had escalated into grappling, the effect of previous wins and losses was no longer apparent and body size strongly predicted contest outcome. Thus, once males can directly assess their opponent's fighting ability, recent fighting experience did not alter the effect of body size on contest behaviour and outcome.     

Male signaling behavior and sexual selection in a wolf spider (Araneae: Lycosidae): a test for dual functions

Male signaling behaviors are often studied in a single context but may serve multiple functions (e.g., in male–male competition and female mate choice). We examined the issue of dual function male signals in a wolf spider species Schizocosa ocreata (Hentz) that displays the same species-specific signaling behaviors in both male–male and male–female contexts. These signaling behaviors have been described as either aggression or courtship according to the context observed. We tested the possibility of dual functions by comparing the relationship between behaviors and outcome of male–male contests (winner/loser) and male–female mating encounters (mating success). Frequency, rate, and mean duration of signaling behaviors did not vary with outcome of male–male contests, which appears instead to be based upon relative size and body mass. Winners of contests had significantly greater body mass than losers, and greater mass relative to opponents was significantly associated with probability of winning. Overall, signaling rates were much higher in male–female interactions than in male–male contests and were higher for males that successfully mated than for those that did not mate. Mean duration of some male displays was also greater for males that successfully mated. However, male size was not associated with probability of mating. Taken together, results suggest an intersexual selection context for the current function of male signals in these wolf spiders and that increased display vigor is associated with male mating success.     

Dominance and reproduction in a parthenogenetic lizard

Summary In most vertebrates, males and females are believed to differ in terms of their investment in offspring. Dominance theory suggests that one way individuals of the sex with lower parental investment can increase reproductive success would be to dominate others of the same sex. The dominant competitors are thought to achieve preferred access to mates, and thus, have greater reproductive success than subordinates.Reproduction in parthenogenetic Cnemidophorus uniparens normally proceeds without males, but individuals exhibit mounting behaviors in captivity that are typical of closely related Cnemidophorus species that reproduce sexually. Thus, these animals provide an unusual opportunity to study the effects of behavior on reproduction apart from copulation and fertilization. In this study relationships between dominance and reproduction were investigated in the unisexual lizard species, C. uniparens. Dominance hierarchies were rapidly established and maintained in the laboratory by agonistic encounters among individuals. The number of times an individual charged its cagemates was positively correlated with the number of clutches and eggs laid. Also, dominant animals who charged their cagemates were likely to win agonistic encounters; recipients of charges usually fled. Hierarchies based on different behaviors were not all related to reproduction. Charges as a predictor of dominance was unrelated to body length, percent increase in body length and time spent in the basking site. However, individuals with a high percent increase in body length spent more time basking. This is likely a result of the increased energy demands of growth in addition to reproduction. Physiological stress as measured by plasma corticosterone titers was unrelated to dominance. We suggest that dominance is an important factor affecting reproduction in C. uniparens.     

Chemosensory cues allow male lizards<Emphasis Type="Italic"> Psammodromus algirus</Emphasis> to override visual concealment of sexual identity by satellite males

Males of many species of lizards show conspicuous breeding colors but, in some species, young competitively inferior males conceal their sexual identity by a female-like dull coloration that allows them to evade aggression from dominant males and to adopt an alternative satellite-sneaking mating tactic. However, large males of the lizard Psammodromus algirus reacted aggressively to young intruder males despite their female-like coloration, suggesting that they might have the ability to recognize competitor males by chemosensory cues. We experimentally manipulated the head coloration (brown vs orange) and scents (male vs female) of small young males. For staged agonistic encounters, we compared the response of resident unmanipulated large males to the different manipulated small males. When we manipulated only the color of small males, the response of resident large males was independent of the paint manipulation; brown and orange males elicited a similar aggressive response. However, when we also manipulated the scent, small males painted orange or brown, but bearing the scent of males, received a significantly higher number of aggressive responses than small males painted orange or brown, but bearing the scent of females. The results showed that, at close range, the reaction of large males to manipulated individuals was dependent on the scent, whereas color seemed to be less important. Coloration may be, however, more important in long-distance communication as shown by the outcome of the first encounters. Also, orange coloration may increase the intensity of the aggressive response. Effective sex recognition by territorial large males is important in natural situations to avoid sneak matings by young male competitors. Thus, even if small males visually conceal their sexual identity, chemosensory cues allow large males to identify them at close range.     

Aggression and fighting ability are correlated in the swordtail fish Xiphophorus cortezi: the advantage of being barless

Males of the swordtail fish Xiphophorus cortezi are polymorphic for the pigment pattern vertical bars. Previous studies indicate that barred males exhibit higher levels of aggression towards males with bars than those without, while barless males fail to exhibit differential levels of aggression to either morph. In this study I matched barred and barless males for size and paired them in dyadic contests in order to determine if either morph was more dominant and if so, if dominance was the result of higher aggression levels. I found that barless males had higher bite frequencies and were able to win a majority of the contests while barred males consistently escalated to biting first, even though in most cases they were ultimately the losers. In order to determine whether the observed aggression levels and fighting abilities were inherent to being barless or a consequence of responses to the bars themselves, the dyads were re-paired once after barless males were given temporary bars and once after barred males had their bars removed. Thus, each morph encountered his opponent in both a barred and barless state. Regardless of bar state, naturally barless males continued to be more aggressive and more dominant than their barred counterparts. In addition, naturally barred males only won contests in which they bit more. These results indicate that for this species, aggression is an important component of winning contests when opponents are roughly the same size. As a result, naturally barless males as a whole appear to have higher resource holding potential (RHP) than naturally barred males of the same size because of their greater aggression levels.     

Rival recognition affects male contest behavior in sand lizards (Lacerta agilis)

Individual male sand lizards meet repeatedly during mate search and engage in costly interactions. If males can recognize rivals, the number of costly fights with a predictable outcome may be reduced. In staged contests between males, second interactions are on average significantly shorter than first ones, suggesting individual recognition. When aggressive behavior differs substantially between the sexes, the more aggressive sex may be predicted to have more variation in skin traits located on the parts of the body displayed during contests; this would facilitate individual recognition. Male (aggressive) sand lizards are more variable in two of three lateral skin traits displayed during males' contests, while females (non-aggressive) are more variable in one dorsal skin trait; in four other dorsal traits there was no difference in variability between the sexes.     

Polyandry and paternity in a wild population of the swordtail Xiphophorus nigrensis

Genetic analyses of parentage provide crucial information about the prevalence of polyandry and the potential for sexual selection to operate in wild populations. In the swordtail Xiphophorus nigrensis, large males are thought to have a competitive advantage due to their superiority in male–male contests and attractiveness to females, who are presumed to mate multiply. I examined the distribution of paternity within broods, the relationship between male body size and paternity and the effect of sire number on fecundity from females collected in the field. Sixty-one percent of females produced offspring from two to four males, with 70% of the offspring typically sired by one of the males represented in the brood. Male body size did not affect paternity share or whether females were multiply mated, as predicted if precopulatory sexual selection has a strong effect on the outcome of postcopulatory sexual selection. Female fecundity increased with the number of sires; however, this relationship was not observed when the smallest broods, where multiple mating is more difficult to detect, were excluded from the analysis. The high levels of multiple paternity and reproductive skew suggest that postcopulatory sexual selection has important evolutionary consequences in X. nigrensis. Traits important in precopulatory sexual selection, such as male body size, however, are more likely to affect sexual selection by increasing the number of mates obtained rather than paternity share within broods.     

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.