Why some rails have white tails: the evolution of white undertail plumage and anti-predator signaling

Conspicuous plumage patches have evolved in birds as conspecific signals for mate attraction and assessment, intersexual competition or to signal alarm. Signals may alternatively be directed at potential predators to discourage pursuit. Rails (Family Rallidae) are ground-dwelling birds, many of which inhabit wetlands, while others occur in forests and grasslands. They are renown for their secretive nature and the tendency to flick their tails when observed. This behavior is more conspicuous in species with white undertail coverts that contrast sharply with darker body plumage. Using species comparisons and controlling for phylogeny, we investigated four hypotheses for the evolution of white undertail coverts in rails. We found little support for the hypothesis that white tails are sexually selected: white tails were not more common in species with polygamous as opposed to monogamous mating systems, species with sexual dimorphism, nor species that display their tails in courtship. Nor did our results support the hypothesis that white tail plumage evolved for intersexual competition during territorial interactions. Instead, we found that species that flock for at least part of the year and species found in open as opposed to concealing habitats were significantly more likely to have white undertail coverts. Rail species inhabiting concealing habitats are less commonly gregarious and more likely selected for crypsis. Using phylogenetically-controlled statistical inference we found that adaptation to open wetland habitats significantly precedes the evolution of white undertails, whereas gregariousness likely evolved later in some lineages. The inferred order of trait evolution suggests that this plumage characteristic could have been selected primarily for enhancement of an anti-predator signal rather than a social signal for conspecifics.     

Serotonin and octopamine elicit stereotypical agonistic behaviors in the squat lobster Munida quadrispina (Anomura, Galatheidae)

Serotonin and octopamine have been implicated as modulators of posture and behavior in several crustaceans. Here we characterize the agonistic behaviors of normally interacting squat lobsters Munida quadrispina (Anomura, Galatheidae) and their responses to serotonin and octopamine injected into the ventral hemolymph sinus, in order to evaluate the potential roles of these amines in modulating agonistic behaviors. Normally interacting M. quadrispina do not develop lasting dominance hierarchies, although transient aggressive and submissive displays do occur. Injected serotonin elicits postures and behaviors in isolated individuals similar to those typical of aggressive, normally interacting animals. Injected octopamine can produce postures and behaviors typical of submissive animals, and elicits behaviors which imply a modulatory role for octopamine in tailflipping. The effects of both amines are reversible and dose dependent, and the dose-response curves parallel the normal progression of agonistic interactions. The social behaviors and reactions to injected serotonin and octopamine of M. quadrispina differ from those of lobsters and crayfish, indicating that interspecific differences in neuromodulation of behavior and motor output exist. Such differences have implications for the understanding of aminergic modulation of aggression and the evolution of aminergic modulation in crustaceans. Accepted: 22 June 1997     

Displays of the Honeyeater Manorina melanocephala

Investigated displays of Noisy Miners, Manorina melanocephala, in Australia. This unusual bird lives in colonies and many ♂♂ care for the offspring of each ♀ flight displays, 11 non-flight displays, and several components of facial displays (including a variable eye patch) are described. The eye patch provides a large yellow and black augmented eye, important in intimidation. No stereotyped sequence of courtship behaviour precedes copulation. Displays are used to advertise nest locations. A greeting display, the corrohoree, is extremely common. The possibility of the evolution of submissive display from threat is discussed. Special vocalisations of ♂♂ and ♀♀ are use in a duet. The maintenance of bonds among many individuals in a colony may be more important than strong pair bonds. Group cohesion is probably maintained by flight display, nest display, mobbing, and other communal activities. High interspecific aggression results in few resident species in colonies. This level of interspecific aggression might be maintained by incorporating much intraspecific mimetic display and ritualised submissive behaviour.     

Interference with ritualized threat behaviour in Cichlasoma nigrofasciatum

According to ethological theory, ritualized threat behaviour functions to prevent overt aggression. This prediction was tested in twenty-three pairs of male Cichalasoma nigrofasciatum. Pairs prevented from giving frontal displays by surgically restrained gill covers showed significantly less mouth fighting than sham operates. Pairs prevented from giving frontals and impaired in delivering tail beats because of separated caudal-fin rays showed intermediate aggressive behaviour. Experimental animals in both groups fought and frequently became dominant when paired with sham operates. It was concluded that threat behaviour in C. nigrofasciatum increases subsequent aggressive behaviours both quantitatively and qualitatively. It is suggested that ethological theory has identified the effects of ritualized aggression as a complex behavioural sequence ending in submissive behaviour with that of its first phase alone.     

Manipulating the appearance of a badge of status causes changes in true badge expression

Signals of dominance and fighting ability (i.e. status signals) are found in a wide range of taxa and are used to settle disputes between competitive rivals. Most previous research has considered status-signal phenotype as an attribute of the individual; however, it is more likely that signal expression is an emergent property that also incorporates aspects of the social environment. Furthermore, because an individual''s signal phenotype is likely to influence its social interactions, the relationships between status signals, social environment and individual quality are probably much more complex than previously appreciated. Here, we explore the dynamic relationship between social interactions and signal expression in a previously undescribed status signal, the frontal shield of the pukeko (Porphyrio porphyrio melanotus: Aves). We demonstrate that frontal shield size is a strong predictor of dominance status within social groups, even after controlling for potentially confounding variables. Then, we evaluate the relationship between social interactions and signal expression by testing whether manipulating apparent shield size influences (i) dominance interactions and (ii) future signal expression. By showing that decreasing apparent shield size causes both an increase in the amount of aggression received and a decrease in an individual''s true shield size, we provide the first evidence of dynamic feedback between signal expression and social interactions. Our study provides important insight into the role of receiver-dependent (i.e. social) costs in maintaining signal honesty and demonstrates a unique approach to studying status signalling applicable to future studies on dynamic morphological signals.     

Impaired social recognition in male mice with repeated experience of aggression

Social recognition is crucial for many aspects of animal behavior in stabilized population. Preliminary data proposed impairment of social recognition in male mice with long experience of aggression. To check this hypothesis, experiments with male mice with different aggressive experience (during 2 and 20 days) were performed. Two types of losers were used as partners: losers with active defense reactions and losers displaying submissive postures. The enhanced aggressive motivation was found in both groups of aggressors. Mice with short aggressive experience demonstrated intensive attacks toward the active losers and decreased aggression directed to submissive losers. Mice with long aggressive experience did not change their behavior depending on a type of the partner and displayed a high level of aggression as a result of dominant aggressive motivation and impaired social recognition.     

Does beak coloration of male blackbirds play a role in intra and/or intersexual selection?

In many bird species, males may show brightly coloured traits and variance in male mating success may be explained by female preference and/or competition between males favouring the most coloured males. Male beak coloration has been suggested to play an important role in the pairing pattern of European blackbirds. Here, we investigate female preference and male-male interactions in relation to male beak coloration in this species. We used a field experiment to measure female and male responses toward stuffed decoys showing either of two beak coloration representing the extremes of the natural variation from yellow to orange. Decoys were situated on the centre of plots and behaviours of males and females approaching the decoy were recorded. The total number of males and females approaching the decoy did not differ between the two model treatments. Similarly, there was no difference in the mean time that males and females spent near the decoy. The number of males displaying aggressive postures did not differ between treatments and no female adopted a sexual posture. We discuss the relevance of our results and potential limitations associated with the experimental procedure.     

Male achromatic wing colouration is related to body condition and female reproductive investment in a dichromatic species,the upland goose

In many bird species, achromatic plumage patch size can serve as a male status signal, but the use of variations in the achromatic colours themselves as a quality signal has only recently come into focus. In our study, we sought to determine whether achromatic plumage reflects individual quality in the upland goose (Chloephaga picta leucoptera). We examined the relationship between male head and wing reflectance, male condition and female reproductive investment. We found that males with darker specula and greater contrast between the white wing coverts and the speculum were in a better body condition. Variations in the brightness of the white plumage were not a quality signal in the upland goose. The information gleaned from the wing colouration of male upland geese could be used during mate selection, when females choose their mate on the basis of the outcomes of aggressive encounters. During these fights, the males expose their white coverts and their specula, which are normally tucked beneath body feathers.     

The strategy of submissive behavior in male mice: the effect of the genotype and the experience of preceding agonistic encounters]

The male mice of two strains with experience of 2 or 10 defeats in intermale agonistic confrontations significantly differ in pattern of submissive behavior (balance of upright and sideways defensive postures, withdrawal, freezing, "on the back" posture). In mice with experience of 20 defeats genetic differences have not been found. The acquisition of consequent experience of defeats does not change the pattern of CBA mice submissive behavior, but significantly increases the share of immobile submissive postures in behavior of C57BL mice. Among submissive males of C57BL strain animals with more active strategy of behavior keep capability for aggressive response to weaker partner. The influence of genotype and previous social contact experience on formation of adaptive in experimental situation strategy of submissive behavior is discussed.     

2. WADERS,DIKKOPS, COURSERS AND COMMON LARKS AT OUDTSHOORN

Eggleton, P. &; Siegfried, W. R. 1979. Displays of the Jackass Penguin. Ostrich 50:139-167.

The paper describes the displays of the Jackass Penguin Spheniscus demersus. It also covers aspects of the species' comfort behaviour. Observations were made at several islands off the South African west coast from 1973 to 1976, although most were made on 130 individually-marked adults at Bird Island (Lambert's Bay) in 1975.

The Jackass Penguin has an extensive repertoire of aggressive displays including Point, Gape, Alternate Stare, Sideways Stare, Peck and Beak Slapping. Other displays are Vibratory Head Shaking (VHS), Ecstatic Display, Mutual Ecstatic Display, several bowing displays and displays associated with copulation. The functions of the displays are discussed and comparison is made with displays in other penguins as documented in the literature. In general the displays of the Jackass Penguin resemble those of other penguins, excluding the genus Aptenodytes. There are, however, variations commensurate with the environment and habits of S. demersus.     

Territorial behaviour in corn mice, <Emphasis Type="Italic">Calomys musculinus</Emphasis> (Muridae: Sigmodontinae), with regard to mating system

We studied territorial behaviour of Calomys musculinus, a promiscuous-polygynous species, in order to test the predictions that during the breeding period, home-range-owner females are more aggressive than home-range intruders and that males do not display territorial behaviour. At the core of the breeding season, we conducted 136 intrasexual and 66 intersexual encounters. The study was carried out in four 0.25-ha enclosures, each containing one independent population. We studied territorial behaviour utilising focal animal sampling to record behaviour. Encounters were conducted in a circular opaque arena (COA) near the home-range’s activity mathematical centre (AMC). Amicable behaviour was not observed in any of the 78 trials among females. The rates of aggressive and submissive behaviour per female–female encounters varied in relation to whether they were home-range owners or intruders, the former being aggressive, the latter submissive. So we could link these drives to the territorial behaviour of C. musculinus. This result fits well with the already known territoriality of C. musculinus during the breeding season. In contrast, the 58 intrasexual male encounters did not show direct aggressive interactions between them. The rates of the different behaviours between males did not vary in relation to whether they were home-range owners or intruders. In our study, which took place during the breeding period, C. musculinus males were nonterritorial, and they never exhibited aggressiveness towards females.     

Prey Escape Direction is Influenced by the Pivoting Displays of Flush-Pursuing Birds

Painted redstart, Myioborus pictus, and its congeners in Central and South America, belong to a small fraction of insectivorous flush‐pursuing birds. Unlike most of the small insectivorous birds, which glean prey from substrates, the flush pursuers spread and pivot their conspicuously patterned tails and wings. This display triggers prey escape flights which are hypothesized to occur through visual stimulation of prey escape circuits [giant descending neuron cluster (GDNC) in Diptera] sensitive to the looming motion of an approaching bird, translational motion of a pivoting body with widely spread tail and contrast of the white‐black plumage pattern. In this paper, data from field observations of redstarts and experiments with bird models show an increase in the frequency of prey escapes away from the strong visual stimulation of an open tail, and in the direction opposite to that of the horizontal translational motion present in the pivots. We discuss how the effect on prey escape direction may enhance prey interception capabilities of redstarts during aerial pursuits. Combined with an earlier study the results show that, unlike the movements of typical gleaner–foragers, the flush displays by redstarts affect prey escape direction in a manner that may facilitate prey tracking and capture by birds. Because the GDNs, which mediate escape initiation, are not sensitive to motion direction, we hypothesize that other neurons, in addition to the GDNs, are involved in influencing the direction of escape responses.     

The neurochemical control of aggression and submission]

Neurochemical mechanisms of agonistic behaviour in different models of aggression are discussed. The effects of aggression and submission experience in 10 mice intermale confrontations under conditions of sensory contact on the levels of brain neurotransmitters and their metabolites were investigated in 7 brain areas. The values obtained in aggressive and control, or submissive and control, animals were compared. In this comparison neurochemical alterations specific for aggressive or submissive behaviours, or nonspecific became apparent. The long experience of victories leads to activation of dopaminergic system through DA catabolism which leads to DOPAC formation. The long experience of defeats increases the 5HT metabolism and decreases NA level in some brain areas. The dopaminergic system of Nucleus accumbens and midbrain are nonspecifically activated in both aggressive and submissive animals. The investigation of values obtained in animals with conversion of behavioural type (after defeat of previously aggressive animals and/or display of aggressive reaction by previously submissive mice) allowed to find many significant differences between aggressive, submissive and "converted" males; in particular the amygdala is the site of opposite changes in 5HT system during inversion of aggressive or submissive behaviours. The above data evidence for the specific role of transmitter systems and brain structures in maintaining or inversion of different types of agonistic behaviour.     

An ethogram of the common marmoset (Calithrix jacchus jacchus): General behavioural repertoire

The behavioural repertoire of four captive breeding pairs of Callithrix jacchus jacchus is described. Social communication took the form of postures, facial expressions, vocalizations and piloerection displays. Detailed analyses were made of piloerection displays, adult play, copulatory, aggressive, and prey-catching behaviour. Aggressive behaviour was uncommon in adult mated pairs. Play between adults showed a degree of temporal organization. Vocalizations were the main methods of intragroup communication whilst piloerection displays were directed towards members of other groups and also to unfamiliar objects. The behavioural repertoire of C. jacchus jacchus is compared with that of other Primates.     

Development and Validation of a Wing Key to Improve Harvest Management of Alcids in the Northwest Atlantic

Abstract: Murres (thick-billed [Uria lomvia] and common [U. aalge]) are legally hunted along the coast of Newfoundland and Labrador, Canada. Razorbills (Alca torda) are also incidentally taken. Only irregular estimates of the total murre harvest are available, so a tool to derive estimates of age- and species-specific harvest is required to effectively monitor the hunt and manage a sustainable harvest. We collected 293 murre and razorbill wings from hunters between 1999–2004, with the goal of identifying wing characteristics that could be used to discriminate age and species. We found that murres and razorbills could be reliably aged (first-yr vs. older) on the basis of molt limits of greater wing coverts. Using a discriminant function (DF) incorporating length of the first primary and second secondary feather, we classified 95–96% of common murres and 99–100% of thick-billed murres correctly to species. First-year thick-billed and common murres also differed in number of pale secondary coverts (median = 12 and 3, respectively), providing another species-specific trait. We developed a key to age and assign species based on these results. We assessed applicability and accuracy of the wing-key with novice observers, who differentiated between murre and razorbill wings using feather-pattern coloration with high accuracy (95 ± 9%) and were able to differentiate between the 2 murres species using 3 techniques: visual assessment of wing shape (83 ± 14% accuracy), the DF (94 ± 6%), and number of worn secondary coverts for first-year birds only (83 ± 5%). Experience increased success rates of aging and species classification using wing shape and number of worn secondary coverts but not using the DF. Despite differences in measurement accuracy and repeatability among observers, the DF proved to be robust. Our results will facilitate implementation of a species composition survey for the murre hunt and will improve identification rates of carcasses found during beached bird surveys in the Northwest Atlantic, aiding in monitoring of alcid populations vulnerable to anthropogenic activities.     

Evidence for the Deflective Function of Eyespots in Wild Junonia evarete Cramer (Lepidoptera,Nymphalidae)

Junonia evarete Cramer is a fast-flying butterfly that perches on the ground with wings opened exhibiting four eyespots close to wing borders. These eyespots presumably function either to intimidate predators, like insectivorous birds, or to deflect bird attacks to less vital parts of the body. We assessed the form, frequency, and location of beak marks on the wings of wild butterflies in central Brazil during two not consecutive years. We found that almost 50% of males and 80% of females bore signals of predator attacks (wing tears), most of them consisting of partially or totally V-shaped forms apparently produced by birds. Males were significantly less attacked and showed a lower proportion of attacks on eyespots than females, suggesting they are better to escape bird attacks. In contrast, females were heavily attacked on eyespots. Eyespot tears in females were higher (and significant different) than expected by chance, indicating that birds do attempt to reach the eyespots when striking on these butterflies. Other comparisons involving the proportion of tears directed or not directed to eyespots in males and females are presented and discussed.     

Role of the pituitary-adrenocortical axis in the control of agonistic behaviour after single and repeated footshock in castrated male mice

Two experiments were conducted to examine the role of the pituitary-adrenocortical axis in the mediation of the effects of single and repeated electric footshock on subsequent agonistic responding in castrated male mice. It was found that (1) preventing corticosterone responses to shock occludes the facilitatory effects of single shock on both aggressive and submissive behaviour and occludes the additional increases in submissive behaviour which normally occur after repeated shock, and (2) blocking pituitary release of ACTH by dexamethasone treatment restores aggressive behaviour after repeated shock, independently of the initial levels of corticosterone and testosterone. These findings suggest that (1) increases in aggressive and submissive behaviour in response to single shock depend on shock-induced increases in corticosterone levels; (2) further increases in submissive behaviour in response to repeated shock require further increases in the corticosterone levels; and (3) inhibition of aggressive behaviour after shock repetition appears to depend on increases in ACTH levels, and occurs independently of the initial corticosterone and testosterone levels.     

Age-related plumage differences of Dunlins along the East Asian-Australasian Flyway

ABSTRACT.   For some populations of Dunlins ( Calidris alpina ), determining the age of individuals on the nonbreeding grounds can be difficult. This difficulty arises in part because some populations undergo their primary molt during the boreal summer, leaving adult and first-year Dunlins with similar amounts of abrasion on their primaries. Ageing Dunlins is further complicated by the presence of adults with buff-fringed, inner median coverts in some populations, a feature often used to age juvenile Dunlins. We examined a number of characteristics helpful in the ageing Dunlins at nonbreeding areas along the East Asian-Australasian Flyway, including: (1) the pattern of white fringes on the inner primary coverts, (2) the color of fringes on the inner median wing coverts, (3) the color of the tip on the carpal covert, (4) the presence or absence of a dark subterminal band on tertial or tertial coverts, and (5) remnants of alternate or juvenile plumage on the belly. Our results demonstrate that the pattern of white fringes on the inner primary coverts is an important character for ageing Dunlin along the East Asian-Australasian Flyway. In addition, we found that all characteristics used for ageing fade with time, and that breeding adults along the East Asian-Australasian Flyway are more difficult to age than adults elsewhere, in part due to the presence of "adult buff" coverts (ABCs) and in part because of the early timing of primary molt. Finally, we suggest that the presence of ABCs might be useful for differentiating the two Dunlin subspecies ( C. a. arcticola and C. a. pacifica ) occurring in Alaska.     

Involvement of dopaminergic systems in the functional specialization of brain areas during formation of aggressive and submissive behavior in mice

In addition to its common activating action, the DA system is involved in the functional specialization of the brain areas which participate in the expression of discrete behavioral components. The evidence for different levels of activity of the mesocortical DA system in aggressive and submissive mice were obtained. In C57BL/6J mice, confrontations produced simultaneous increase in the extracellular DA content and its release from the nerve terminals in the nucleus accumbens and frontal cortex, while an elevation of the HVA concentration in these structures was found only in submissive mice. After 20 encounters, the habituation of animals to the repeated stress exposures and conditioning developed. Activation of the DA metabolism (increase in the DOPAC level and DOPAC/DA ratio) in the hippocampus was observed only in aggressive mice after 20 days of confrontation, when the extinction of the information novelty leading to aggression had been already accomplished. Our findings suggest the predominance of the role of the mesolimbic DA system, in particular, of its mesoaccumbens link, in the extinction of the information novelty in aggressors. A role of the mesocortical DA system in realization of the submissive behavior patterns, stress reactions, conditioned defensive behavior, anxiety-related behavior, and in modulation of the anxiety response to social stimuli is considered.     

Escape behaviour in blue‐winged grasshoppers,Oedipoda caerulescens

Blue‐winged grasshoppers Oedipoda caerulescens (Linnaeus, 1758) are commonly found in flat, open, unprotected areas. In the event of immediate danger, they leave their camouflaged position and jump away at the last moment. The present study conducted in a flight arena shows that, despite jumping at short notice from a crouching position, the grasshoppers achieve the correct timing for an optimal leap. If both compound eyes are blinded and the animals are stimulated by touch to execute an unprepared jump, the take‐off of the flightless nymphs is delayed, and adults are delayed in raising their wings; the animals tumble backward during the leap (in the case of adults, if they do not open their wings). This is a result of the unprepared take‐off position; because the entire length of the hind legs cannot be used for acceleration, the body is rotated backward. However, the escape path is not ultimately affected because, in the air, physical processes compensate for the unfavourable starting conditions. In addition, no disadvantage is evident upon landing. In each case, a hook landing was completed safely (i.e. the grasshopper landed and swung round to face the direction it had come from). The impact force is reduced and the grasshopper stabilizes itself by rotating from a forward to a backward position, immediately after the first contact with the ground. The hook landing also serves to confuse the potential attacker, and the disappearance of the bright blue hind wings of the adult makes it difficult for predators to shift quickly enough to a different kind of search to relocate their prey. In conclusion, the present study shows that the escape behaviour of blue‐winged grasshoppers is adapted to extremely short escape distances.