Transgressive aggression in Sceloporus hybrids confers fitness through advantages in male agonistic encounters

1.  We investigated agonistic behaviour and associated characteristics of Sceloporus woodi (Florida scrub lizard), Sceloporus undulatus (Eastern fence lizard) and their hybrids using staged territorial encounters.
2.  These Sceloporus hybrids exhibit transgressive aggression and transgressive head-girth relative to the parental species and the transgressive aggression was specifically associated with an advantage in agonistic encounters. Our results suggest a hybrid advantage in natural habitats when defending and invading territories against either parental species.
3.  We further analysed general advantages in agonistic encounters across the entire three-group system to elucidate characteristics that may be advantageous under specific circumstances. Individuals with larger body size (SVL) and greater aggression had an overall advantage in agonistic encounters; however, smaller individuals could win when slightly more aggressive and fatter, and less aggressive individuals could win when slightly larger, especially with greater head-girth.
4.  The extreme hybrid phenotypes likely occurred through transgressive segregation, which has been implicated as a process through which homoploid, hybrid speciation can occur. Some form of ecological divergence is necessary, however, to impede parental gene flow. Our data suggest that ecological divergence could manifest in territorial species through transgressive aggression.     

Intra- and interspecific agonistic behavior of the subterranean termite Microcerotermes crassus (Isoptera: Termitidae)

The aim of our study was to investigate the intra- and interspecific agonistic behaviors exhibited by the worker and soldier castes of the subterranean termite Microcerotermes crassus Snyder (Isoptera: Termitidae). Aggression between M. crassus colonies from different field locations and also against three termite species--Coptotermes gestroi (Wasmann), Globitermnes sulphureus Haviland, and Odontotermes sp.--were observed in the laboratory. Termite responses were tested in paired combination of castes (soldiers versus soldiers, soldiers versus workers, and workers versus workers) consisting of 10 individuals each. Significant agonistic behaviors were observed only in encounters between pairings of different termite species. M. crassus was aggressive toward individuals from different species but not toward individuals from different M. crassus colonies. Mortality of M. crassus reached 100% in most of the interspecific encounters. However, no or low mortality was recorded in the intraspecific pairings.     

Influences on Vigilance in Nine-banded Armadillos

Functional aspects of vigilance in nine-banded armadillos (Edentata: Dasypus novemcinctus) were investigated. Data on solitary individuals revealed few age or sex differences in time spent vigilant. However, vigilance increased in most social contexts and was highest during intraspecific agonistic encounters. These results suggest social functions for armadillo vigilance. Armadillos with higher vigilance levels had longer flight distances, suggesting that vigilance may also have antipredator benefits. Further examination of flight distances revealed that they varied with age, weather conditions, and time of year (breeding versus non-breeding season), and were negatively correlated with body size. In total, the results indicate the complexity of vigilance in a solitary species and suggest that armadillo vigilance is multifunctional.     

Body size as a factor determining dominance in staged agonistic encounters between male brown anoles (Anolis sagrei)

The results of staged agonistic encounters between males indicate that body size is an important determinant of dominance in male brown anoles (Anolis sagrei). Larger males defended their perch sites more successfully than did smaller males, perched higher than smaller males, and were most often the first male to enter the other male's territory. Larger males also exhibited more challenge displays than did smaller males. Head-nods, a display given by subordinate individuals, were observed only in smaller males.     

Agonistic interactions with asymmetric body size in two adult-age groups of the annual killifish Millerichthys robustus (Miller & Hubbs, 1974)

In this study, the author evaluated two adult age groups of the Mexican rivulus Millerichthys robustus with body size asymmetries to determine the strategies used by an annual killifish during agonistic interactions of different ontogenetic stages. To achieve this goal, the author first characterized the ethogram of agonistic interactions of M. robustus composed of seven behavioural units in males and five behavioural units in females. The author then analysed agonistic interaction strategies used by males and females with body size asymmetries in two groups of different adult ages that represent different ontogenetic stages: (a) just after sexual maturity was reached, at 5 weeks of age, and (b) near natural death, at 24 weeks of age. The agonistic behaviour patterns of M. robustus were compatible with the logic of mutual assessment. Large males had an advantage during their interactions in both age groups, winning all of the encounters. Nonetheless, there was more aggression in 5-week-old fish encounters. In addition, small 24-week-old fish were more aggressive than small 5-week-old fish. These changing strategies may be because of the cost–benefits required during a fight at each ontogenetic stage. In the female encounters, size did not predict winners, as both small and large fish won a similar number of encounters, and some contests remained unresolved regardless of age group. There was a tendency for small females of any age to risk more than males in fights to maintain reproductive fitness.     

Previous Experiences Alter the Outcome of Aggressive Interactions Between Males in the Crayfish, Procambarus Clarkii

Various measures of size have been associated with increased likelihood of winning such agonistic encounters between decapod crustaceans. Recent social experience, in the form of 'winner' or 'loser' effects, has also been shown to alter the outcome of encounters between size-matched individuals. This study further explores the importance of social experience in crustacean agonistic encounters. Crayfish, Procambarus clarkii , of different size classes were given a series of winning or losing experiences to determine whether this would affect their likelihood of success in subsequent fights. Recent wins or losses in a series of fights did alter the likelihood of an individual winning a subsequent encounter, despite an apparent disparity in size between opponents that would otherwise serve as an accurate predictor of the outcome.     

雄性大仓鼠体重对其斗殴行为及社会等级的作用

在实验室条件下测定雄性大仓鼠体重对社会等级和斗殴行为的作用模式,检验体重对雄性大仓鼠社会等级及斗殴行为序列具有重要影响的假设。本实验以16只成年雄性大仓鼠为目标个体,采用等级内部的线性概率、组内循环三元组数量（d）和优势等级的线性度（K）,排列个体的社会等级序位。研究结果表明,雄性大仓鼠可形成近似线性的优势等级,体重与个体的优势等级,攻击行为和胁腺标记行为均呈显著的正相关关系,与防御行为和攻击潜伏期存在显著的负相关关系。说明独居性物种大仓鼠雄体间可形成优势等级关系,体重对此关系具有重要的作用。     

Intraspecific variation in structures that display competitive ability: large animals invest relatively more

The benefits of possessing (or enlarging) a display structure used in competitor assessment will vary between individuals of a species. For some individuals the cost of developing such a feature could outweigh any benefits accrued. Animals whose competitive ability is high can benefit from display, but there is little advantage in advertising low competitive ability. Thus, since large animals generally win fights it can be predicted that large animals should have relatively large display features. This prediction was tested using data on the moorhen, Gallinula chloropus. Evidence is presented that in this species frontal shields are used in competitor assessment. The size of an individual's frontal shield varied seasonally. Maximum shield size corresponded with peak aggressive behaviour. Shield size was positively correlated with body weight, which is the best predictor of the outcome of agonistic encounters in this species, and the slope of the regression was significantly greater than one. Thus shield size proved to be positively allometric as predicted. It remains to be tested whether or not positive allometry is a feature of all structures used to display competitive ability.     

Levels of social tolerance between snow volesChionomys nivalis during over-wintering periods

We examined winter social behaviour of the snow voleChionomys nivalis Martins, 1842, a rock-dwelling microtine found principally in high-mountain biotopes. We conducted dyadic encounters among wild-caughtC. nivalis to determine patterns of intraspecific tolerance and the degree of aggressiveness between and within sexes during wintering periods. Our data show a high frequency of agonistic behaviours in all social encounters. Male pairs spent less time being aggressive than female pairs but more time in investigative behaviours. In contrast, the intensity of aggression displayed during interactions was significantly higher in male-male encounters than in female-female encounters, with an intermediate level of aggression in male-female encounters. The low degree of social tolerance found seems to be in agreement with the available data on the social organization of this species and further suggests that individuals may not spatially associate during over-wintering periods. Although both males and females were socially intolerant, the different patterns observed between the sexes may underlie different strategies.     

Morphological traits determine the winner of “symmetric” fights in hermit crabs

This study aimed at investigating in the hermit crab Pagurus longicarpus whether some morphological traits and their slight variations might determine the winner of a contest in symmetric pairs, i.e. pairs composed of individuals matched by their overall size. In the pre-experimental phase, 400 crabs were individually kept in isolation for a week to eliminate the effects of their previous social experiences and were allowed to enter adequate shells to equalize their motivation to fight. Then, we formed 200 pairs matched for shell and body size and observed their agonistic behaviour for 15 min. Alphas (and betas) were deemed as those individuals that won more than half of the recorded fights. Finally, crabs were sacrificed and sexed, and 6 and 8 measures were taken of their shells and bodies, respectively. Raw data were converted to compositional data and subjected to an isometric log-ratio transformation prior to statistical analysis. Alphas were found to occupy heavier and wider shells than betas, both characteristics that, together with the correlated high level of encrustation, make the apparent dimension of the crab larger and thus favour that individual during agonistic encounters. As expected, hermit crabs were sexually dimorphic in their chelae, but sexes differed also in the length of the dactylus of their third walking leg and in their body weight. Alphas were heavier than betas and had longer propodi and wider dactyli on the third walking legs than betas. The latter character might be advantageous during shell fights when the walking legs of the attacker are used to hold the defender's shell in the spasmodic shaking and their wider dactyli might assure a firmer seizing of it. Indeed, a correct execution of this pattern allows for the eviction of the defender. Our results suggest that more attention towards variations in morphological traits other than body size might improve our understanding of the factors that affect individual Resource Holding Power and fight dynamics.     

Aggression and sexual activity of male Southern elephant seals, Mirounga leonina

The agonistic and sexual activity of individual male Southern elephant seals (Mirounga leonina) was studied during the breeding season at South Georgia. Harem bulls were older and larger than non-harem bulls and, although size and age are related, harem bulls were also larger than non-harem bulls of the same age. Large body size conferred advantages in fighting and the agonistic relationships of the bulls gave rise to a dominance hierarchy with access to harems and activity within harems, being determined by rank. Vocalizations were the most highly rank-related aggressive behaviours. The most common vocalization, the roar, probably functioned in both individual recognition and size assessment. Over 90% of encounters of high-ranked bulls were "walkovers" and only 4% of 4003 agonistic interactions nvolved physical contact.     

Species-specific differences in dynamics of agonistic interactions may contribute to the competitive advantage of the invasive signal crayfish (Pacifastacus leniusculus) over the native narrow-clawed crayfish (Astacus leptodactylus)

Species-specific differences in dynamics of agonistic interactions may influence the outcome of interspecific competition and potentially contribute to competitive advantage of one species over another. In this study, we compared the dynamics of agonistic interactions of one of the most successful crayfish invaders of European freshwater ecosystems, the signal crayfish (Pacifastacus leniusculus) and the widespread native European species currently undergoing range expansion in Croatia and Europe, the narrow-clawed crayfish (Astacus leptodactylus). Comparisons between P. leniusculus pairs and the A. leptodactylus pairs demonstrated significant differences in frequency and duration of agonistic encounters: P. leniusculus dyads engaged in fewer fights, but their duration was significantly longer. In staged interactions between size-matched interspecific pairs, agonistic behaviour of P. leniusculus individuals translated into dominance over their A. leptodactylus counterparts. This indicates that the success of P. leniusculus in agonistic encounters with the native competitor might stem from its readiness to continue fighting for a longer time period, and could lead to ecological advantages during niche competition even when facing a successful native crayfish species.     

Seasonal changes in and effects of familiarity on agonistic behaviors of rat-like hamsters (Cricetulus triton)

The seasonal changes in agonistic behaviors and effects of familiarity on agonistic behaviors in wild-caught adult rat-like hamsters (Cricetulus triton) were observed in dyadic encounters in a neutral arena. The aggression of opposite- and same-sex encounters became higher or remained the same during the non-breeding season. This indicates that the hamsters were solitary during both seasons. Familiarity increased the aggression in male–male encounters and decreased the aggression in female–female encounters during both seasons. Familiarity also increased the aggression in female–male encounters during the non-breeding season and had no effect on the aggression in female–male encounters during the breeding season. These results may be related to the hamsters social structure. The more agonistic acts both male and female hamsters had, the more frequently they marked using flank glands during both seasons. This implies that flank gland marking can be used to advertise status and can be assessed by opponents to reduce the agonistic costs.     

Influence of male morphology on male mating status and behavior during interunit encounters in western lowland gorillas

The western lowland gorilla (Gorilla gorilla gorilla) is one of the most sexually dimorphic primate species. Mature males are twice the size of females and have grey fur on their backs and a fibrous, adipose crest on their heads. Such traits are likely to have evolved by sexual selection, either because they confer advantages during male-male fights or because females prefer males with more dimorphic traits. We developed photogrammetric methods for distance collection of morphological data from silverback males frequenting the Lokoué forest clearing in Odzala-Kokoua National Park, Republic of the Congo. Body length, head-crest size, musculature development, and extent of the grey color on the back were assessed in 87 nonbreeding and breeding mature males. Behavioral data were also collected during 312 male-male encounters involving 67 mature males in order to estimate their level of aggressiveness. The number of females belonging to a mature male positively correlated with the male crest size, body length, and musculature. Whereas morphological variables did not significantly affect the intensity of male-male encounters, the number of females attending male-male encounters strongly affected the number of agonistic displays by the two males. We discuss the mechanisms through which males with more exaggerated traits could obtain a mating advantage, namely male-male fights or female mate choice.     

Significance of Agonistic Vocalization in the Croaking Gourami (Trichopsis vittatus,Teleostei)

Trichopsis vittatus emits high amplitude sounds during agonistic encounters with conspecifics. The sound producing organ is derived from the structural components of the pectoral fins. The study involved muting a sample of subjects by removing two pectoral fin tendons without any further restriction in movements and behaviour. Mute and unaltered males were then placed together in pairs and the following agonistic behavioural elements were determined: attacks, lateral displays, sound production and frontal displays. Soniferous males had a higher probability of winning contests when size differences were small. In pairs with big size ratios, the larger males were more often successful. Besides visual and acoustical signals, lateral line stimuli seem to play no role in threatening displays. These results demonstrate that vocalization during agonistic encounters is important for becoming dominant in specific circumstances. Because of correlation between main frequencies and size, sound emission might be a very effective method of assessing the physical strength of an opponent.     

Intra- and Inter-colonial Agonistic Behavior in the Termite, Microcerotermes fuscotibialis Sjostedt (Isoptera: Termitidae: Termitinae)

Pairing termite workers from cultures obtained from same colony of Microcerotermes fuscotibialis Sjostedt did not result in agonistic behavior or mortality of the paired individuals. Termite pairings from different colonies located at different distances resulted in serious encounters and aggressive behavior leading to significant mortality of the individuals. Subtle agonistic behavior was observed when colony mates were reunited after months of separation, but this did not result in any significant mortality. These results confirmed observations of non-overlap of foraging trails of these termites in the field which averts possible encounters and conflict. Therefore these termites are able to recognize their colony mates irrespective of period of separation, a factor for colony affiliation, kinship and recognition.     

Individual recognition in crayfish (Cherax dispar): the roles of strength and experience in deciding aggressive encounters

The outcomes of agonistic interactions modulate access to resources and thereby affect fitness. Success in agonistic encounters may depend on intrinsic physical and physiological performance, and on social experience. Here we test the hypothesis that previous experience will override physical strength in determining the outcome of fights in the freshwater crayfish Cherax dispar. Between unfamiliar opponents, greater chelae closing force significantly increases the chances of winning. However, even when the chelae of the original winners were disabled, the winners kept on winning against the same opponents after 30min and 24h. This winner effect disappeared when previous winners encountered unfamiliar individuals. Similarly, a previous loss did not affect the outcomes of subsequent encounters with unknown crayfish. We suggest that this prolonged recognition of individuals and their relative fighting ability is a mechanism that can reduce the number of agonistic encounters experienced by individuals.     

Agonistic interactions between nymphs of Lycorma delicatula (Hemiptera: Fulgoridae)

The nymphs of Lycorma delicatula typically aggregate on tree branches for feeding. We studied agonistic interactions between nymphs in the field and investigated the effects of prior residence and body size on contest outcome. Resident nymphs maintained positions on tree branches and engaged in feeding, whereas intruding nymphs moved along tree branches. When an intruder approached, the resident typically raised its front legs as a threat posture. The intruder would either move away or palpate the resident, and the resident often lowered its body when palpated. The intruder would then attempt to shake the resident back and forth, with the number of attempts varying from one to several. The resident either maintained its position or fled. After a contest, the winner was determined to be the one occupying the contested position on the branch. The resident nymphs won 84.2% of the agonistic encounters, and the nymphs with larger body size won 63.2% of the agonistic encounters. Logistic regression analysis revealed that prior residence was a significant factor for the contest outcome, but body size was not. We discuss the hypotheses for the dominance of the residents in agonistic interactions between nymphs of L. delicatula.     

Coral reef fish assemblages: intra- and interspecific competition for shelter sites

Synopsis Observations were made on intra- and interspecific aggressive interactions among the fishes living in the rubble/sand coral reef habitat in St. Croix, U. S. Virgin Islands. Four species (beaugregory — Stegastes leucostictus; ocean surgeonfish — Acanthurus bahianus; doctor fish — A. chirurgus; common squirrelfish — Holocentrus rufus) which sheltered in holes on the reef all actively defended one to several shelter sites at dusk. Short-term shelter side fidelity was observed in three of these four species. Agonistic interactions over both food and shelter occurred during the daytime but much less frequently than agonistic interactions over shelter at dusk. Dominance in intraspecific aggression was determined almost completely by the relative sizes of the individuals involved, with the larger individuals dominating in 95–98% of all encounters. A similar, but less strong, relationship between size and dominance existed for interactions between closely related species. For aggressive encounters between unrelated species, however, both relative sizes and species identity determined the outcome. Species, both diurnal and nocturnal, which strongly defend several shelter sites may have a strong and disproportionate impact on the sheltering behavior of other fishes. Intraspecific and interspecific defense of shelter sites may determine the patterns of mortality that result from predation, thereby influencing population abundances and assemblage composition.