Anti-Predator Signals as Advertisements: Evidence in White-Throated Magpie-Jays

Calls and displays elicited by predators usually function as alarms or to inform predators of their detection. However, predator encounters may afford some individuals the opportunity to demonstrate quality or signal their availability. Here, I report on a class of vocal signals produced in predator-elicited displays that share many characteristics with sexually selected song. White-throated magpie-jays ( Calocitta formosa ) display at low-threat predators while producing 'loud display calls' (LDCs). I use this term because the calls occur primarily in two display contexts (see below) though occasionally in other contexts as well. Such calls and displays are primarily produced by males, and also occur in one other context, at dawn. Playback experiments showed that despite being elicited by predators, males were more likely than females to respond to LDCs, and more likely to respond when their mate was fertile. Over 134 different call types were produced in over 200 displays by 34 males; the largest minimum repertoire size was 67. Presentations of taxidermic raptor mounts elicited some LDCs, but fewer calls and lower diversity than at dawn or in predator approach displays. The male bias and high diversity suggest that LDCs are an outcome of intersexual selection, while their elicitation by predators suggests an alarm function. I propose that male magpie-jays use predator encounters as opportunities to advertise their presence and availability as mates; they use LDCs as songs. Such a communication system seems to have been favored by the unusual social system of magpie-jays, in which female groups defend territories and males have little opportunity to defend resources for mate attraction, forcing them to advertise when females are paying the most attention, during predator encounters.     

Characterization of Australian fur seal vocalizations during the breeding season

The vocal repertoire, structure, and behavioral context of airborne vocalizations produced by Australian fur seals (Arctocephalus pusillus doriferus) are described using recordings made at a breeding colony on Kanowna Island, Bass Strait, Australia. The study identified six different call types: three produced by males (bark, guttural threat, and submissive call); five produced by females (bark, guttural threat, submissive call, growl, and pup attraction call) and the female attraction call produced by pups and yearlings. Vocalizations were compared according to age and sex classes. The overall structure and function of the pup attraction and female attraction call produced by females, yearlings, and pups, was similar. However, while similar in their overall appearance, certain call types have a lower fundamental frequency when compared with other fur seals. In addition, the male bark call alters in rate of production according to the context used, where calls are slower when males are stationary and advertising their territorial status and faster when males are involved in confrontations with other males or actively herding females. Further research is required to investigate changes in environmental conditions and their effects on shaping the call structure and communication in Australian fur seals.     

Vocal behavior of captive Sichuan golden monkeys (Rhinopithecus r. roxellana)

Vocal behavior of four pairs of adult Sichuan golden monkeys was studied at the San Diego, Seattle, Los Angeles, and San Francisco zoos between November 1984 and October 1986. Four call types made up 86% (1,357/ 1,578) of vocalizations that occurred during 45 h of tape-recording. All four major vocalization types were made by both sexes. There was, however, significant sexual asymmetry in their production: 82% to 99% of each type was produced by one sex or the other. Frequently, duets developed when an individual male or female responded to the vocalizations of its cagemate. Males vocalized more than did females in each of the four pairs, and males responded vocally to female calls more than twice as frequently.     

Singing in the Face of Danger: the Anomalous Type II Vocalization of the Splendid Fairy-Wren

Males of certain species of fairy-wrens (Aves: Maluridae) emit a unique vocalization, the Type II vocalization, in response to the calls of potential predators. We conducted field observations and playback experiments to identify the contexts in which the Type II vocalization is emitted by splendid fairy-wren ( Malurus splendens ) males, and to examine social and genetic factors that influence its occurrence. In field observations and controlled playback experiments, Type II vocalizations were elicited most consistently by calls of the predatory gray butcherbird ( Cracticus torquatus ). Some vocalizations from other avian species also elicited Type II vocalizations, and the majority of these were vocalizations from avian predators. Splendid fairy-wrens are cooperative breeders, and males that responded with Type II vocalizations to playbacks of butcherbird calls tended to be primary rather than secondary males, had larger cloacal protuberances, and were older than those that did not respond. In addition, secondary males that were sons of resident females were more likely than non-sons to respond with a Type II vocalization. In another playback experiment, females responded similarly to the Type I song and Type II vocalizations of their mates. Although the Type II vocalization is emitted primarily in response to predator calls, it is inconsistent with an alarm call explanation. Patterns of reproductive success among Type II calling males suggest that it does not function as an honest signal of male quality. At present, the function of the vocalization remains anomalous, but indirect fitness benefits may play a role in its explanation.     

Conflicting Territory Use in Males and Females of a Monogamous Ungulate,the Kirk's Dikdik

In monogamous mammals it is often unclear why males do not defend larger territories to attract more than one female. I investigated the territoriality of the monogamous Kirk's dikdik, Madoqua kirki, a dwarf antelope, in which food resources increase with territory size and some males defend enough resources for more than one female. Yet, all males are paired monogamously. When males were removed from small territories, their female partners spent more time outside of their territories than females in large ones. When females were removed, their male partners almost never left. Pairs in small territories spent more time together than pairs in large ones. Paired males left mostly together with their females, apparently not on their own initiative. Presumably because females in small territories left more often, their males spent more time outside in the female's company than males in large territories. I argue that males in smaller territories can keep better track of their females and that they can effectively reduce their females' time outside. Male intrusion pressure was unrelated to territory size, but it increased in the presence of unguarded females. If large territories decrease the ability to mate guard, and if unguarded females attract competing males, then defending large territories may be uneconomical, even it they could attract more than one female. On the other hand, territories must be large enough to satisfy the requirements of a single female.     

Food-associated calls in rhesus macaques (Macaca mulatta): I. Socioecological factors

Upon discovering food, free-living rhesus macaques (Macaca mulatta)on the island of Cayo Santiago, Puerto Rico, produce a complexof vocal signals consisting of five acoustically distinguishablecalls. This report examines the socioecological factors elicitingcall production and the information protentially conveyed toothers. The primary contexts for three vocalizations ("warbles," "harmonicarches, " and "chirps") are encounters with rareand highly preferred foods (e.g., coconut). Two other vocalizations("coos" and "grunts") are produced both in food (primarily provisionedchow) and in nonfood contexts, such as during mother-infantseparation and grooming interactions. Grunts given upon encounteringfood are acoustically distinct from those given in nonfood contexts.In contrast, coos associated with food are statistically indistinguishablefrom coos given in other contexts. When conspecitics hear thesefood-associated calls, they typically approach the caller. Coosare less likely to lead to approach than other food-associatedcalls, Results from all-day follows on adult males and adultfemales reveal that changes in hunger level influence call ratebut not call type; the different call types are produced throughoutthe day. We infer that the structure of food-associated callsprovides information about the quality of the food discovered,whereas call rate conveys information about the relative hungerlevel of the caller. In this population, adult males give fewerfood-associated calls than adult females. In addition, femaleswithin large matrilines call more than females within smallermatrilines, and males who are resident in a group are more vocalthan peripheral males.     

Long-range call use in dominance-structured Crested Tit Parus cristatus winter groups

In this study on free-ranging Crested Tits Parus cristatus , I examine the relationship between social dominance and the frequency of use of long-range communication calls. Calling rates of trills were highest among socially dominant individuals and they gave more calls when close to the boundary of their territories. Dominant females uttered fewer calls than their mates. However, they gave significantly more calls than subordinate males and subordinate females, the latter calling least. A removal and playback experiment revealed a relationship between the utterance of trilled calls and the defence of the winter territory in the Crested Tit. Although territorial vocalizations could incur costs, territorial individuals may gain from improved winter survival by decreasing the risk of food stealing by Crested Tits from adjacent territories.     

Ultrasonic Vocalizations in Rat Sexual Behavior

Ultrasonic vocalizations are very conspicuous during rat matingactivity. Two types of calls are produced by both sexes. Thefirst, brief complex calls with the main frequency centeredabout 50 kHz, occur primarily in conjunction with solicitationand mounting activity. The second type of call is the long,22 kHz whistle which is emitted mainly by the male during thepostejaculatory refractory period, but also by both male andfemale at other times during the copulatory sequence. The occurrenceof ultrasonic vocalizations is correlated with sexual motivationof rats. Males emit more 50 kHz calls before successful matingtests than before tests in which they fail to ejaculate. Furthermore,more vocalizations are emitted by the pair prior to intromissionsthan prior to mounts without intromission. Just before ejaculationthere is a large increase in the rate of calling and, at times,transition by the male to calling at 22 kHz. This latter eventmay represent physiological dearousal by the male. Followingejaculation, the male characteristically emits 22 kHz vocalizationsand exhibits a sleep-like EEG pattern. The function of the postejaculatoryvocalization may be to enforce separation between the matingpair, while at the same lime maintaining contact between thepartners. Fifty kHz calls, on the other hand, prime and facilitatesexual responsiveness of the female. Tape recorded vocalizationsof mating rats facilitate solicitation behavior of estrous femalesin the presence of castrated males, and such females also showa preference for these sounds in a "Y" maze. Deafening of femalesdoes not affect their normal pacing of copulatory contacts,but it drastically reduces their solicitation behavior. Thestudies summarized in this paper lead us to conclude that ultrasonicvocalizations play a major role in the integration of reproductiveactivity in the Norway rat, Rattus norvegicus.     

Food, audience and sex effects on pinyon jay (Gymnorhinus cyanocephalus) communication

Pinyon jays (Gymnorhinus cyanocephalus) have a complex social system that may require a complex communication system. They need to interact with multiple flock members, and they form life-long pair-bonds. We researched whether pinyon jays would selectively vocalize depending on the presence or absence of food and certain flock members. We recorded the vocalizations of nine pinyon jays (four pair-bonds and one single male) in response to different audience types. The calls of the test bird were recorded after it was given either an empty food cup or one containing 50 pinyon pine (Pinus edulis) seeds, and the bird was in the presence of one of the following audience types: (1) two males and two females including subject's mate; (2) two males and two females excluding subject's mate; (3) four males excluding mate; (4) three females excluding mate; and (5) no audience. Birds gave fewer calls when there was food. When alone, birds called in a manner that may maximize long-distance transmission. Trends indicate that birds call differently to their mate. A sex effect was also found in that males and females called in a distinct manner, possibly reflecting differences in dominance status. Overall, birds responded to the presence or absence of an audience.     

Correlation between male size and territory quality: consequence of male competition or predation susceptibility?

Territory characteristics correlate with male characteristics in several species. This can result from male competition for the best territories, or from males varying in their ability to pay other costs of territoriality, such as predation risk costs. In a population of threespine sticklebacks, Gasterosteus aculeatus , we found the biggest males to defend the biggest territories with a low structural complexity and a high female encounter rate. By experimentally manipulating competition intensity and habitat structure, we show that both male competition and predation exposure influenced the distribution of territories among males. Males increased the size of their territory when a neighbouring male was removed, whereas they reduced their territory when habitat complexity and cover from predators were reduced, with large males reducing their territory size less than smaller males. This suggests that large males occupy large, open territories both because of their superior competitive ability and because of their either lower predation susceptibility or higher risk-taking. Large, open territories were beneficial in mate attraction and male competition and predation exposure therefore biased mating opportunities towards large males. This suggests that cost of territoriality to males may reduce mate choice costs to females by securing that large males are encountered more often than small males, and by providing an additional cue, territory quality, which indicates which males are worth inspecting.     

Significance of temporal and spectral acoustic cues for sexual recognition in Xenopus laevis

As in many anurans, males of the totally aquatic species, Xenopus laevis, advertise their sexual receptivity using vocalizations. Unusually for anurans, X. laevis females also advertise producing a fertility call that results in courtship duets between partners. Although all X. laevis calls consist of repetitive click trains, male and female calls exhibit sex-specific acoustic features that might convey sexual identity. We tested the significance of the carrier frequency and the temporal pattern of calls using underwater playback experiments in which modified calls were used to evoke vocal responses in males. Since males respond differently to male and female calls, the modification of a key component of sexual identity in calls should change the male's response. We found that a female-like slow call rhythm triggers more vocal activity than a male-like fast rhythm. A call containing both a female-like temporal pattern and a female-like carrier frequency elicits higher levels of courtship display than either feature alone. In contrast, a male-like temporal pattern is sufficient to trigger typical male-male encounter vocalizations regardless of spectral cues. Thus, our evidence supports a role for temporal acoustic cues in sexual identity recognition and for spectral acoustic cues in conveying female attractiveness in X. laevis.     

Perception of female reproductive state from vocal cues in a mammal species

While acoustic signalling by males is known to affect male-male competition, mate attraction and the timing of ovulation, the extent to which sexual selection has shaped the evolution of female acoustic signals is poorly understood. Among mammals, experimental evidence indicates that females attract mating partners by using olfactory and visual signals to advertise their reproductive state. Whether or not males ascertain female reproductive state from vocal signals has, however, never been systematically tested. In this study, we use playbacks of recorded vocalizations to demonstrate that male Barbary macaques, Macaca sylvanus, can discriminate between female copulation calls given at different stages of the oestrous cycle, responding more strongly to those given around the time when conception is most likely to occur. Acoustic analysis suggests that the mean dominant frequency of call units and a number of temporal parameters could provide males with the information necessary to discern the proximity of ovulation in this way Our results provide the first experimental evidence that the calls of female mammals may contain information on reproductive state, which males can perceive and use in such a way as to increase their reproductive success.     

Mate-locating behavior of the butterfly Lethe diana (Lepidoptera: Satyridae): do males diurnally or seasonally change their mating strategy?

The mate-locating behavior of male butterflies has been classified into two major types, territorial and patrolling. Territorial males defend a specific site, whereas patrolling males fly around a wider area without having to defend a site. In this study, I investigated the use of these tactics by males of the satyrine butterfly, Lethe diana. A previous study suggested that the males of L. diana change their mate-locating behavior during the day (they patrol in the morning and defend territories in the afternoon) and that patrolling is the primary mating strategy, whereas defending territories is a supplementary one. In the present study, I found that the daily activity pattern of the males of L. diana was similar to that described in the previous study: males often flew around in the morning and competed for territories in the afternoon. However, contrary to the previous study, all courtships and copulations were performed within male territories during their territorial activity. Closer observations revealed that copulations found in male territories were achieved by the owner of the territory. Males tended to feed in the morning, suggesting that the males flying in the morning searched for food rather than females. I conclude that territory holding is the primary male matelocating tactic in L. diana. I further found that, in summer, males exhibited territorial behavior later than in spring or autumn, which may be a strategy for preventing heat stress.     

Recognition of family-specific calls in stripe-backed wrens

Males of the cooperatively breeding stripe-backed wren Campylorhynchus nuchalis, learn repertoires of stereotyped calls (termed WAY calls) from older male relatives. As a result, these vocalizations are normally specific to patrilineal family groups but are sometimes shared by male relatives in different groups. To determine whether or not this species can recognize the calls of different family groups, I performed playback experiments with individual call types recorded from males of known social and genealogical relationships. Subjects discriminated between the calls of unrelated neighbouring groups and unfamiliar groups, and they discriminated both of these from calls of their own groups. However, subjects failed to distinguish calls of males in other groups from calls of their own groups when these males were members of the same patriline. These results indicate that stripe-backed wrens can discriminate between repertoires of these calls that match or differ from their own. Consequently, they can recognize members of their patriline, not just members of their immediate group. These vocalizations probably provide a useful mechanism for recognizing group membership in this species and might also provide a mechanism for recognizing unfamiliar relatives in other groups. Copyright 1999 The Association for the Study of Animal Behaviour.     

人为疼痛刺激下金色中仓鼠幼鼠叫声的发育过程

金色中仓鼠幼鼠可以利用可听声和超声信号与母鼠进行交流,这些声音可以反映个体的寒冷、饥饿、疼痛等不同生理状态和需求。因疼痛诱发的叫声对于维持幼鼠的存活也有着重要意义。本实验通过录制人为疼痛刺激下,不同日龄金色中仓鼠的叫声,分析并比较了5 - 30 日龄金色中仓鼠不同发声信号特征(可听声出现频次、可听声持续时程、可听声主频率、超声出现频次、超声持续时程、超声主频率)的性别差异;同时记录了幼鼠的两种发声随日龄的变化趋势。结果发现:幼鼠两种叫声的各种参数均未表现性二型现象。幼鼠超声发生频次随日龄增加而逐渐减少。而可听声却正相反,随日龄增加而逐渐上升,并在20 日龄时达到最高峰,之后逐渐下降。可听声持续时程也随日龄而增加。综上,在人为疼痛刺激下,随着日龄增加,幼鼠更倾向于使用可听 声而非超声与母兽进行交流。     

The Haremic Mating System and Mate Choice in the Wide-Eyed Flounder, Bothus podas

The social structure and reproductive behaviour of the wide-eyed flounder, Bothus podas, was studied in the coastal waters around the Azorean Islands. Both sexes are territorial throughout the year. Adult males defend large territories, which include several smaller female territories. Intraspecific agonistic behaviour was frequent and differed between sexes: males were more aggressive towards other males, while females were only aggressive towards each other and juveniles. During the reproductive season and only at dawn, territorial males court and mate successively with females in their territories, and females seem to show mating fidelity to their dominant male. Such territoriality and mating patterns indicate a haremic social system in the wide-eyed flounder. In order to identify potential factors influencing female mate choice acting on this haremic system, we examined male mating success and some of its potential correlates. We found no evidence for female preference for any of the males' physical or territory characteristics. However, courtship effort was strongly correlated with the total number of attempted and successful spawnings, indicating that females seem to mate preferentially with males that court them more vigorously. Thus, our data suggest that courtship plays an important role in determining male mating success in the wide-eyed flounder and, that it may possibly serve as an honest indicator of male `quality' for female choice.     

The social organization of a sexually dimorphic honeyeater: the Crescent Honeyeater Phylidonyris pyrrhoptera,at Wilsons Promontory,Victoria

The social organization of the sexually dimorphic and dichromatic Crescent Honeyeater Phylidonyris pyrrhoptera (Latham 1801) was studied at Wilsons Promontory National Park between January 1994 and January 1997. All breeding attempts at one site were followed during the spring and summer of 1996. Most pairs were multibrooded, with pair-bonds maintained throughout the study. Breeding males held well-defined territories. Females, although generally restricting their activities to within their mate’s territory, visited other males’ territories more frequently than did their mates and did so close to the time of egg laying. Males aggressively chased intruding males, but appeared to permit intrusions by non-mate females. Circumstantial evidence suggests some males may kill neighbour’s nestlings. Female Crescent Honeyeaters are solely responsible for nest construction and incubation. While both sexes attended nestlings, males contributed to parental care significantly less than females. Males also provided significantly less parental care than has been reported for monomorphic species of honeyeater. Single syllable calls are given throughout the year. Peaks in the rate of double and multisyllable calls corresponded with peaks in breeding activity. Only males were observed uttering loud multisyllable calls. In contrast to predictions arising from sexual selection theory and observations of the mating system of sexually dimorphic hummingbirds, the population of Crescent Honeyeaters studied exhibited a socially monogamous mating system. However, the very limited male role in parental care and non-observance of male territory boundaries by females suggest the genetic mating system may not be one of monogamy.     

Acoustic Monitoring Reveals Congruent Patterns of Territorial Singing Behaviour in Male and Female Tropical Wrens

Many animals defend territories against conspecific individuals using acoustic signals. In birds, male vocalizations are known to play a critical role in territory defence. Territorial acoustic signals in females have been poorly studied, perhaps because female song is uncommon in north‐temperate ecosystems. In this study, we compare male vs. female territorial singing behaviour in Neotropical rufous‐and‐white wrens Thryothorus rufalbus, a species where both sexes produce solo songs and often coordinate their songs in vocal duets. We recorded free‐living birds in Costa Rica using an eight‐microphone Acoustic Location System capable of passively triangulating the position of animals based on their vocalizations. We recorded 17 pairs of birds for 2–4 consecutive mornings and calculated the territory of each individual as a 95% fixed kernel estimate around their song posts. We compared territories calculated around male vs. female song posts, including separate analyses of solo vs. duet song posts. These spatial analyses of singing behaviour reveal that males and females use similarly sized territories with more than 60% overlap between breeding partners. Territories calculated based on solo vs. duet song posts were of similar size and similar degrees of overlap. Solos and duets were performed at similar distances from the nest for both sexes. Overall, male and female rufous‐and‐white wrens exhibit very similar spatial territorial singing behaviour, demonstrating congruent patterns of male and female territoriality.     

Effects of androgens on social behavior and morphology of alternative reproductive males of the Azorean rock-pool blenny

In the Azorean rock-pool blenny Parablennius sanguinolentus parvicornis two sequential reproductive tactics occur. Larger and older males establish breeding territories, while some of the smaller males become attached to nest-holder territories, acting as satellites on these territories, which they help to defend while trying parasitic fertilizations when females go in the nests to spawn. In the present paper we tested the effects of the androgens 17alpha-methyltestosterone (MT) and 11-ketotestosterone (KT) in the expression of male secondary sex characters and bourgeois behavior in satellite males. One week after satellites were implanted with Silastic tubes containing MT, KT, or castor oil (control), androgen-treated satellites had developed male secondary sex traits such as longer and wider male-type genital papilla and anal glands that secrete a sex pheromone, both traits being less expressed or absent, respectively, in satellite males. Androgen treatment had no effect on the gonadosomatic index or on the development of the testicular gland. KT treatment had a positive effect on relative liver weight. In terms of behavior, androgen-implanted individuals were less aggressive both in a mirror test and toward females when these were introduced into their tanks. MT-treated individuals spend more time inside the provided nests. Only androgen-implanted satellites managed to have the females entering their nests. When given a chance in a group tank either to try to attract females to their own nest or to act as satellites of an already established nest-holder's nest, MT-implanted males spent significantly more time in their own nest than near the nest-holder nest. These data suggest that androgens, particularly testosterone, may be involved in mating tactic switching in this species.     

The Role of Learning in the Aggressive and Reproductive Behavior of Blue Gouramis, Trichogaster trichopterus

Like countless other vertebrates and even many invertebrates, blue gouramis, Trichogaster trichopterus, a freshwater tropical fish, can learn to associate environmental cues with the appearance of biologically important events. Research with blue gouramis reveals that this capacity for learning, which psychologists call Pavlovian conditioning, provides the means to enhance their territorial defense as well as reap large reproductive benefits. That is, males are able to defend their territories more efficiently when the appearance of a rival is signaled than when territorial invasion is less predictable. Not only does signaling enable Pavlovian-conditioned males to mount a more aggressive defense of their territories than males that do not have the benefit of signaling, but it helps them to concentrate their efforts at times when, or places where, the territory is most vulnerable. In addition, both males and females can learn to anticipate the appearance of a mate. Signaling of female accessibility enables Pavlovian-conditioned males to attenuate their initial aggressive response to arriving females without compromising their territorial defense. More important, when the arrival of a potential mate is signaled, Pavlovian-conditioned males are able to spawn with females sooner, clasp females more often, and produce more young than males that do not have the benefit of a signal. The ability of blue gouramis, among many other fish, to anticipate the appearance of such different biologically important events suggests that Pavlovian conditioning may play a significant role in their behavioral ecology. In addition, conditioning may provide a useful tool in several practical domains, including animal husbandry, zoological management, and conservation biology.