The organization of play fighting in the grasshopper mouse (Onychomys leucogaster): Mixing predatory and sociosexual targets and tactics

The body targets contacted, the type of contact made, and the patterns of defense and counterattack elicited by those attacks are examined in the play fighting of captive male and female pairs of grasshopper mice. The nape was the most frequently contacted body target, irrespective of the type of contact made, be it nosing, allogrooming, biting, or striking with a forepaw. The types of defense varied with both body area contacted and type of attack performed. Based on the topography and pattern of contact, it was concluded that grasshopper mice, as is the case for many other muroid rodents, primarily attack and defend targets otherwise contacted during precopulatory encounters. However, grasshopper mice, which are obligate carnivores, also attack and defend predatory targets, although less frequently than sociosexual targets. Surprisingly, predatory attacks were more likely to be counterattacked with predatory attacks, whereas sociosexual attacks were more likely to be counterattacked with sociosexual attacks. Conspecific aggression involves bites directed at the face, lower flanks, and dorsum. Neither the biting of these areas nor the tactics of attack and defense usually associated with such bites were observed during the juvenile interactions. There were no sex differences in either frequency or patterns of attack and defense in play fighting. The data presented for grasshopper mice shed light on the issue of mixing behavior patterns from multiple functional systems during play. Aggr. Behav. 26:319–334, 2000. © 2000 Wiley‐Liss, Inc.     

Play-fighting differs from serious fighting in both target of attack and tactics of fighting in the laboratory rat Rattus norvegicus

Play-fighting is often difficult to differentiate from inhibited or immature serious fighting because both may utilize many of the same behavior patterns. In the rat the two behaviors involve different targets of attack. During play-fighting, snout or oral contact is directed at the opponent's nape of the neck, whereas during serious fighting, male residents mostly direct their bites at the intruder's rump. Although similar to those used in serious fighting, the behavior patterns used during play-fighting are modified to achieve the different targets of attack. Even though the tactics of attack and defense appear more adult-like with increasing age, the playful targets persist well into adulthood.     

Agonistic versus amicable targets of attack and defense: Consequences for the origin,function, and descriptive classification of play-fighting

Play-fighting appears to involve the behavior patterns of attack and defense otherwise seen in serious fighting. The degree of similarity, however, depends on the body targets attacked and defended during these forms of fighting. For many taxa, including diverse mammalian families and some birds, the same targets are attacked and defended during both play-fighting and serious fighting. However, for several species of muroid rodents, the targets of play-fighting are not the same as those of serious fighting. In these cases, the tactics of attack and defense are also different. It is argued that for these muroid species the playful targets have arisen from amicable behavior (e.g., social investigation, greeting, allogrooming) rather than, as appears to be the case in so many other taxa, from agonistic behavior. These data strongly suggest that “play-fighting” has evolved from different precursors in different taxa and thus has multiple origins. Furthermore, these data have an important bearing on the universal applicability of many of the suggested functions of play-fighting and also on how such behavior is to be described and classified.     

Targets and tactics: The analysis of moment-to-moment decision making in animal combat

Even though injury and death are more common consequences of fighting among animals than once believed, they are still relatively infrequent. Modern evolutionary models of animal combat have emphasized that given the threat of retaliation, animals only escalate to more injurious fighting if the benefits outweigh the costs, and then only if threat and bluff fail to achieve the goal. Such models stress the role of communication as to whether animals decide to escalate or not. An alternative view is that failure to produce injury or death arises from the neutralization of one animal's attack by another's defense. That is, attack and defense end in a stalemate that may be misinterpreted by outside observers as an absence of injury producing behavior. As attack typically involves the biting or striking of specific body targets, movements and postures occurring during combat need to be analyzed with respect to their role in gaining or averting such contact. For example, in the combat of muroid rodents the attacker targets the lower dorsum and flanks (low threshold) or face (high threshold), whereas a defender may defensively launch counterstrikes against the attacker's face. Two combat tactics (supine defense and lateral attack) typically present in the fighting of muroid rodents are analyzed in detail to illustrate how targets constrain the movements of combatants. Such a functional analysis of combat assumes that the movements and postures performed are related to their role in the attack and defense of targets. Deviations from such a strict functional interpretation reveal some of the other factors that may constrain the combatants' behavior. For example, body morphology and the aggressiveness of the opponent are shown to be important in deciding the type of combat tactic to use and how it is performed. Finally, movements and postures that are neutral or even counterproductive for attack and defense may be revealed as communicatory. This approach provides a means of analyzing behavior during the "heat of combat" that is typically not dealt with in traditional evolutionary models. Aggr. Behav. 23:107–129, 1997.© 1997 Wiley-Liss, Inc.     

Relationship between attack and feeding in the insect predatory behavior of grasshopper mice

In grasshopper mice, the consequences of feeding and state of hunger affect the predatory attack to a limited extent. The purpose of this study was: (1) to establish other ways that attack and feeding were connected besides the two effects already mentioned; and (2) to determine if caching of dead prey could account for the limited connection between attack and feeding. Experiment 1 showed that attack and feeding were connected as part of a reaction chain. Experiment 2 showed that both shared a common maturational process despite the fact that each response achieved an improved efficiency in a different way. Experiment 3 showed that caching prey was not correlated with the killing of a prey and thus could not account for the limited connection between attack and feeding. It is proposed that a highly canalized attack response specifically buffered against the effects of feeding and hunger plays an important role in the mouse's carnivorous life style.     

Identification of the possible origin of the body target that differentiates play fighting from serious fighting in syrian golden hamsters (Mesocricetus auratus)

Play fighting in the Syrian Golden hamster Mesocricetus auratus can be distinguished from serious fighting by the targets attacked in each case. In play fighting, the animals attack and defend the cheeks and cheek pouches, whereas in serious fighting they attack and defend the rump and lower flanks. Since play typically involves the use of behaviors borrowed from other functional contexts, this paper investigates the origin of the cheek target during play fighting. Comparison of resident-intruder serious fighting with awake and anesthetized intruders does not reveal the cheek to be an inhibited target for serious attack. Similarly, analysis of social investigation and allog-rooming, while revealing the ears to be important targets, do not show the cheeks to be targets in these behaviors. Sniffing, licking, and nibbling of the cheek area appear to occur mainly during sexual encounters by males. This area, seemingly a sexual target, may be the one utilized during play fighting.     

Targets of attack and defense in play-fighting of the Djungarian hamster Phodopus campbelli: Links to fighting and sex

Analysis of the body targets attacked and defended during play-fighting by juvenile Djungarian hamsters Phodopus campbelli revealed that about 70% of all attacks were directed at the mouth. If successfully contacted, the mouth was briefly licked and nuzzled. The remaining playful attacks were gentle bites directed at the rump, and to a lesser extent, the top of the head. During serious fighting the top of the head and the rump are targets of attack, whereas the mouth is not. Licking and nuzzling the mouth was found to be a behavior performed by adult males at the beginning of sexual encounters. Therefore, play-fighting in juvenile hamsters cannot be thought of merely as a form of “mock fighting” since the principal target is seemingly sexual, not agonistic. The data also show that of the sexual body targets contacted, adult females are more likely to defend the mouth. In this way it is suggested that targets attacked and defended during juvenile play-fighting are derived from adult contexts in which such targets are defended. This hypothesis accounts for the prevalence of agonistic targets in the play-fighting of many species, and may provide a rationale for classifying those amicable targets that are competed for during play-fighting.     

Targets and tactics of agonistic and precopulatory behavior in montane and prairie voles: Their relationship to juvenile play-fighting

Play-fighting by juvenile montane and prairie voles involves attack and defense of the head, neck and shoulders. Since during play animals typically borrow behavior patterns from other functional contexts, two adult behavioral contexts were compared to juvenile play-fighting. These were serious fighting and sexual encounters. During serious fighting in a resident-intruder paradigm, most bites are directed at the rump and lower flanks. During sexual encounters, especially in precopulatory behavior, the head, neck and shoulders are gently contacted. Therefore, play-fighting by juveniles would appear to involve attack and defense of areas of the body contacted in adult precopulatory behavior, not adult fighting. Furthermore, the species-specific differences in juvenile play-fighting were also found to be matched by species-specific differences in precopulatory behavior. In both playful and precopulatory encounters, montane voles contacted the head and used upright defensive behaviors more often than prairie voles. In contrast, prairie voles made mutual contact more often and were more likely to rotate to supine in defense of contact to the nape and head. These findings support our hypothesis that juvenile play-fighting in muroid rodents involves the precocial expression of precopulatory, not agonistic behavior.     

Offensive and defensive bite-target topographies in attacks by lactating rats

The attacks by resident lactating Wistar rats on sexually naive conspecifics of both sexes were examined. Male and female intruders were equally attacked in terms of frequency and number of bites, but the topographies of biting seen in these encounters were different. Similarly to male-male agonistic interactions, females were attacked in a fashion which avoided bites to the head and snout (“offensive” attack), whereas males were frequently bitten on such vulnerable regions (“defensive” attack). This dichotomy in bite pattern suggests that different motivations and functions underlay maternal aggression in these situations. The defensive attack on males may be a deterrent to infanticide since only male intruders counterattack lactating females and kill their pups. The attack on females may be concerned with resource competition.     

Supine defense in the intraspecific fighting of male house mice mus domesticus

House mice have been reported rarely to perform the supine behavior pattern as a defensive tactic during intraspecific fighting. However, in this study of intraspecific fighting by male mice, it is shown that mice do indeed rotate to supine. This maneuver is used to evade or extricate themselves from bites to the lower dorsum by the attacking opponent. Once free from the bite the defender does not remain supine, but will immediately turn to prone and flee. Remaining motionless in the supine position may serve a submissive function in other species, but this does not seem to be the case for mice. The present findings illustrate that the supine tactic is a dynamic maneuver for defense of body areas targeted by the opponent. © 1992 Wiley-Liss, Inc.     

Prey-dependent effects of fluprazine hydrochloride on predatory aggression in northern grasshopper mice (Onychomys leucogaster) and rats (Rattus norvegicus)

Treatment with the phenylpiperazine DU 27716 (fluprazine hydrochloride) inhibited the predatory killing of albino mice by northern grasshopper mice (Onychomys leucogaster) and of frogs by rats. This treatment had no effect on cricket predation by grasshopper mice or worm predation by rats. The prey-specific effect of fluprazine did not seem to result from a decreased tendency to attack nor was its effect restricted to prey showing characteristic rodent defensive responses. Rather, the drug seemed to increase fearfulness when the prey exhibited vigorous defensive behavior. It is possible that increased fearfulness induced by treatment with this drug may also contribute to its antioffensive effect during conspecific encounters.     

Infanticide and maternal aggression: Synchrony of male and female reproductive strategies in mice

Thirty-six percent of male mice from three strains attacked newborn pups sired by another male. No male attacked its own offspring. Females did not show differential aggression toward males likely (strangers) or unlikely (sires) to attack their pups. Both forms of aggression were unaffected by housing in rooms which did or did not contain the aggression targets. The three strains differed in strength of maternal aggression but not in the incidence of infanticide. Females showed more aggression when mated with males of the same, rather than a different, strain but no differences with intruders of the same or a different strain. Infanticide by males is best viewed as a postcopulatory, intermale-competition strategy, and maternal aggression as a counter strategy.     

Effects of toxicosis on the predatory behavior of the golden hamster (Mesocricetus auratus)

The purpose of this study was to measure the effects that toxicosis had on the hamster's predatory behavior and to compare these results with those found in previous studies with grasshopper mice. A 0.15 M LiCl injection caused hamsters to develop a greater aversion and to inhibit their feeding and attack responses more frequently toward a house cricket than did a similar injection of NaCl. The added presence of an almond coating on the cricket prolonged the number of days that a hamster exhibited an aversion toward the prey and an inhibition to attack. Essentially, the inhibitory effects from toxicosis in the hamster paralleled those found with the grasshopper mouse. The differences in these inhibitory effects are attributed to inherent differences in the attack responses of the two rodents.     

Importance of tactile and olfactory cues to the inhibition of the grasshopper mouse's attack through toxicosis

This study examined the effect that toxicosis paired with the presence of a distinctive texture had on the inhibition of the grasshopper mouse's predatory attack. The first experiment measured the context in which tactile cues would be most effective by presenting prey with various combinations of added stimuli. The combination of distinctive tactile, olfactory, and gustatory cues produced the longest lasting inhibition. Because certain manipulations also had an unintended weak odor associated with them, the second experiment measured the importance of a weak or strong odor to inhibition of a mouse's attack. A combination of strong odor and distinctive texture paired with toxicosis inhibited an attack more effectively than a weak odor and a similar texture. In a third experiment, toxicosis was paired with an acraea moth caterpillar which has a highly distinctive texture. This produced the longest lasting inhibition of mouse's attack observed thus far. The ecological significance of a combination of a distinctive texture and odor to the inhibition of the grasshopper mouse's attack is discussed.     

Specialised use of working memory by Portia africana,a spider-eating salticid

Using expectancy–violation methods, we investigated the role of working memory in the predatory strategy of Portia africana, a salticid spider from Kenya that preys by preference on other spiders. One of this predator’s tactics is to launch opportunistic leaping attacks on to other spiders in their webs. Focussing on this particular tactic, our experiments began with a test spider on a ramp facing a lure (dead prey spider mounted on a cork disc) that could be reached by leaping. After the test spider faced the lure for 30 s, we blocked the test spider’s view of the lure by lowering an opaque shutter before the spider leapt. When the shutter was raised 90 s later, either the same lure came into view again (control) or a different lure came into view (experimental: different prey type in same orientation or same prey type in different orientation). We recorded attack frequency (number of test spiders that leapt at the lure) and attack latency (time elapsing between shutter being raised and spiders initiating a leap). Attack latencies in control trials were not significantly different from attack latencies in experimental trials, regardless of whether it was prey type or prey orientation that changed in the experimental trials. However, compared with test spiders in the no-change control trials, significantly fewer test spiders leapt when prey type changed. There was no significant effect on attack frequency when prey orientation changed. These findings suggest that this predator represents prey type independently of prey orientation.     

The influence of the victim on shock induced aggression in rats.

In two studies, free-roaming male rats (aggressors) were shocked in the presence of male target rats restrained in either an upright or a supine posure. In addition, in Experiment II, two levels of aggressor shock intensity (0.8 mA or 2.0 mA) were used while targets received one of three levels of shock (0.5 mA, 1.5 mA, or 2.5 mA). In both studies, upright targets were attacked less than supine targets. Frequency of aggression was directly related to level of aggressor shock intensity in Experiment II. Also, attack by 0.8-mA aggressors against supine targets was inversely related to level of target shock intensity. The low level of attack against upright targets was interpreted in terms of a threat diaplay. Similarily, it was concluded that the target shock-intensity effect in Experiment II was due to specific threat behaviors displayed by those supine rats that received the highest-intensity shocks.     

Pictorial target control of schedule-induced attack in White Carneaux pigeons

Three pigeons with a history of attacking a mirror target, and two of six pigeons with no prior exposure to targets, attacked a colored photograph of a conspecific during exposure to intermittent schedules of reinforcement for key pecking. Rate of attack on the photograph decreased when the reinforcement schedule was removed. The topography, temporal pattern, and locus of attack on the picture were comparable to schedule-induced attack on live, stuffed, and mirror targets. When silhouette, outline, and plain paper targets were used, schedule-induced attack was more sensitive to a change in target characteristics with a concurrent target-preference procedure than with an analogous successive-testing procedure. The combined results of the two testing procedures indicated that an “upright” white-on-black silhouette of a pigeon with or without an eye was more effective in controlling attack than was a comparable “inverted” silhouette, an outline of a pigeon, or a piece of colored paper.     

Does fear modulate defensive and offensive types of maternal attack in mice?

Swiss CD‐1 lactating mice show a different pattern of attack toward intruders of differing sex, displaying defensive attack against the male (bites on the head and ventrum associated with fear) and offensive attack against the female (bites on the back and flanks with no elicitation of fear). This dichotomy may reflect diverse functions of maternal aggression: the attack toward males (the more infanticidal gender in laboratory strains) has been interpreted as a counterstrategy to infanticide, whereas the attack toward females may serve to establish a social hierarchy or to space rivals of the same sex. In terms of proximal mechanisms, fear may be a key factor involved in the modulation of the different patterns of attack. In Experiment 1 we compared the pattern of attack of lactating females in Swiss CD‐1 and Wild mice toward male and female intruders in relation to fear components of behavior of the attacking dams. Results showed that in Swiss mice, male intruders were attacked with a defensive type of attack accompanied by high levels of fear, whereas female intruders did not elicit fear in the attacking animal but were attacked with an offensive pattern. In Wild mice, both types of intruders were attacked with a defensive pattern; notwithstanding, fear was evident only toward male intruders. This suggests that fear is not totally responsible for the expression of the defensive type of attack. To test the hypothesis that defensive attack toward male and female intruders may be related to the infanticidal potential of the intruder, Experiment 2 examined levels of infanticide in both male and female Swiss CD‐1 and Wild mice. Swiss female mice showed virtually no infanticidal behavior, whereas Swiss males and both sexes of Wild mice showed similarly high levels of infanticide (55%–75%). From a game theory perspective, the defensive pattern of maternal attack toward female intruders in Wild mice is discussed as “extreme” defense of a high value resource and thus, functionally, a competitive form of aggression. Aggr. Behav. 26:193–203, 2000. © 2000 Wiley‐Liss, Inc.     

Effects of the cholinomimetic drug arecoline upon aggression: Intra- vs. inter-specific allocation of attack

Systemic injections of cholinomimetic drugs have been reported to induce both rage and predatory attack in several species. In order to assess the relative contribution of each of these two behavioral patterns in the control of cholinergically induced attack, a group of adult female cats was chemically stimulated in the simultaneous presence of both a prey object and a conspecific attack object. In this choice situation stimulated cats initially tended to engage in rage attack. When a second group of subjects was tested in a successive choice situation a significantly greater number of attacks occurred against conspecifics. The results suggest that cholmergic stimulation initially induces affective attack, with somewhat less frequent incidents of predation.     

Effects of female presence on intrasexual aggression in male lizards,Podarcis hispanicus

Resource value and expected gain in reproduction may affect motivation to fight and the likelihood of winning. Previous experiments have showed that males increase their fighting effort when defending a territory that contains females. However, we hypothesized that for an intruding lizard, the value of a new territory may be lower if he already has a female in his own territory, and consequently, aggressivity should be lesser than if he has no access to any female. We staged encounters between males of the lizard Podarcis hispanicus in outdoor terraria to analyze the outcome and detailed behaviors involved in agonistic interactions in the presence or absence of a female in the terraria of resident and intruders. Our results showed that when a female was present, the level of aggressivity of the resident male was higher; the probability of winning the contest also increased, but only if the intruding male had no females in his own terraria. In contrast, when the intruding male was also the owner of another territory containing a female, residents were less aggressive. We suggest that the lack of information on the reproductive state of an unfamiliar female may be enough to decrease fighting motivation of an intruding male, if he has more expectations of success with his own familiar female. We conclude that differences in expected reproductive success with different females may help to decide the outcome of conflicts between males quicker and cheaper. Aggr. Behav. 28:491–498, 2002. © 2002 Wiley‐Liss, Inc.