Males are attracted by their own courtship signals

Courting male fiddler crabs Uca terpsichores (1 cm carapace width) sometimes build mounds of sand called hoods at the entrances to their burrows. Males wave their single enlarged claws to attract females to their burrows for mating. It was shown previously that burrows with hoods are more attractive to females and that females visually orient to these structures. In this study, we test whether males also use their hoods to find their burrows. We first determined the maximum distance that males can see and find a burrow opening without a hood. Males were removed from their burrows and placed on the sand at a range of distances from a burrow opening. If they were more than about 8 cm (seven units of eye-height) away, they were unable find the burrow. In contrast, males that were burrow residents used a non-visual path map to return to their burrows from much greater distances. To determine if hoods help males find their burrows when there are errors in their path maps, we moved residents 1–49 cm on sliding platforms producing errors equal to the distances they were moved. Males with self-made hoods or hood models at their burrows relocated their burrows at significantly greater distances than did males with unadorned burrows. Hood builders also relocated their burrows faster. Hence, hoods have two functions: they attract females and they provide a visual cue that males use to find their burrows quickly and reliably when their path maps fail. An erratum to this article can be found at     

Neighbor burrow-plugging in Ilyoplax pusillus (Crustacea: Brachyura: Ocypodidae)

The burrow living ocypodid crab Ilyoplax pusillus (De Haan, 1835) sometimes plugs the burrow of neighbors situated from 1.0 to 8.3 cm away with surface mud, while the neighbor, termed the pluggee, is in the burrow. Most pluggers were large males, whereas pluggees were smaller than pluggers and had a sex ratio close to 1:1. After being plugged, most pluggees usually reemerged on the surface within 5 min, but occasionally took up to 77 min to do so. The plugger always foraged or performed waving displays around the burrow of the pluggee while the pluggee stayed inside the burrow. Although the activity site of the pluggee was originally directed toward the burrow of the plugger or the adjacent area, after reemergence its activity site was oriented toward another direction. From these observations, it is suggested that neighbor burrow-plugging is a behavior adopted mainly by large males as a means of maintaining the area of their surface activities against smaller neighbors. My observations were made from June 1984 to June 1985 at Fukuro River Estuary, central Japan.     

Testing function of fiddler crab claw waving by manipulating social context

Many territorial advertisement signals are thought to be dual-function signals, directed to both rival male and receptive female conspecifics. However, few studies have tested this assumption by examining whether in fact both sexes are likely to elicit signaling behavior from territorial males. In this study, I experimentally manipulated the social context of male sand fiddler crabs (Uca pugilator) to investigate the effect of different audiences on the performance of the claw-waving display, a territorial signal that is often presumed to be directed to both males and females. To test whether males perform this signal to both audiences, I measured the frequency of waving behavior by focal males when housed in field enclosures alone, with only males, with only females, or with both males and females. Focal males waved at a low frequency when alone, and the presence of males had no effect on their level of waving. However, in the presence of females, focal males showed a significantly higher level of waving, whether or not males were also present. In addition, there was no association between fighting and waving behavior. This experiment provides evidence that from the perspective of the signaling male, the claw-waving display of U. pugilator is not a dual-function signal but rather is primarily directed to receptive females. Received: 16 December 1999 / Received in revised form: 1 February 2000 / Accepted: 19 February 2000     

Deterring predators, daunting opponents or drawing partners? Signaling rates across diverse contexts in the lizard Anolis sagrei

Although the visual display behavior in Anolis lizards has received ample attention, the function of dewlap extensions (DE), push-ups (PU), and head-nods (HN) in general, and in Anolis sagrei in particular, remains highly equivocal. Therefore, our primary goal was to quantify the display rates of these visual signal types (DE, PU, and HN) in a variety of contexts, using A. sagrei as study species. To our knowledge, this is the first study to test individuals of both sexes in a repeated-measures design across multiple contexts, including predator, non-predator and social interactions (mirror, male–male, male–female, female–male). We found that males have an overall higher signaling rate than females across all contexts. In addition, we found that lizards of both sexes exhibited higher display rates in the presence of conspecifics than when confronted with a predator or non-predator, suggesting that DE, PU, and HN function in intraspecific communication, not in predator deterrence. Whereas females did not significantly raise display rates in a consexual and heterosexual context with respect to subject-alone context, males did. The PU signal type only appears to play a major role for A. sagrei males during aggressive encounters. During heterosexual interactions, increased frequencies of all signal types suggest that DE, PU, and HN are essential for male courtship. Finally, we suggest that intersexual selection is probably a driving force for frequency-related dewlap use in both sexes. In contrast, pronounced intersexual differences were detected for PU and HN rates within a social context.     

Effect of female molt stage and sex ratio on courtship behavior of the blue crab Callinectes sapidus

In many species, males and females actively participate in courtship, and the outcome of pre-mating interactions influences the mating success of both sexes. Female blue crabs, Callinectes sapidus, mate soon after their final molt to maturity; thus female molt stage dictates the timing of mating. In a field experiment, we manipulated female molt stage and sex ratio to test their effects on the courtship behavior of both sexes, if female behavior influences the behavior and pairing success of males, and if male courtship influences male pairing-success. Early-molt-stage females avoided males during courtship, whereas late-molt-stage females sought out males. As a result, males had to pursue and capture early-molt-stage females whereas males displayed to late-molt-stage females and more easily physically controlled them. Males sometimes abandoned late-molt-stage females, but this occurred more often when females were abundant. The rate at which females avoided males was positively correlated with that of males abandoning females, and males that were unsuccessful at pairing met with higher rates of female resistance than successful males, suggesting that female behavior influences male pairing-success. Unlike unsuccessful males, successful males more often made the transition between display and maintaining physical control of the female. At high male sex ratios, males initiated courtship more readily; thus both sexual competition and female behavior influence male courtship in this species. Received: 7 July 1996 / Accepted: 10 January 1998     

Are vocal signals used to recognize individuals during male–male competition in greater prairie-chickens (Tympanuchus cupido)?

Vocal signaling can be an important component of vertebrate communication during social interactions. If vocalizations vary among individuals but are consistent within a given individual, they may be used to discriminate among individuals. In many species, territorial males use vocalizations to discriminate between neighbors and strangers and either respond more aggressively toward strangers relative to neighbors (“dear enemy” effect) or they respond more aggressively toward neighbors relative to strangers (“nasty neighbor” effect). In the greater prairie-chicken (Tympanuchus cupido), male vocalizations are an integral part of the display males produce on leks. We investigated whether male greater prairie-chickens discriminate among familiar individuals on their own territory, familiar individuals outside their normal territory and strangers from a nearby lek. Vocal characteristics varied among males, suggesting that vocalizations may potentially be used by prairie-chickens to identify individuals. Males responded to playback of prairie-chicken calls by vocalizing at a faster rate and approaching the playback speaker, but did not vary in their response to the vocalizations based on the identity of the caller. Our results suggest that variation is present among the vocalizations of individual male greater prairie-chickens, but males do not appear to discriminate among familiar individuals and strangers based solely on their “boom” vocalizations. Greater prairie-chicken vocalization likely functions as a way of announcing that a territory is occupied and defended, but it may also serve as a way of advertising to conspecifics or as a signal that is secondary to other forms of communication.     

A conspicuous courtship signal in the fiddler crab Uca perplexa: female choice based on display structure

The fiddler crab Uca perplexa has a conspicuous male courtship signal that is directed at females to attract them to the male’s burrow for mating. The signal involves waving the unflexed large claw up and down. To determine whether the spatiotemporal structure of the wave is under selection by female choice, we examined whether females had a preference for any particular features of the wave. Females respond to a waving display by either visiting the male’s burrow entrance or by electing to pass without visiting the burrow. We filmed mate-searching females and the waving males that they visited or passed. We documented the wave structure of these males using frame-by-frame analysis. Males produce a two-part wave with component A preceding component B. Both components have an upstroke, a pause at the apex and a downstroke. The tip of the claw was raised much higher in B than in A. Visited males had a shorter delay between the two wave components than did males that the females passed without visiting. Visited males also produced component B waves that had a slower upstroke than those of passed males. There was a significant correlation between the relative height of the raised claw and the duration of the upstroke of component B. Females were selecting males that raised their major claw to the highest position (two to three times as high as the carapace width). Passed males brought down their major claw earlier and from a lower position than did visited males. The data suggests that wave structure has evolved through female choice. Male display rate and body size were not female choice cues. An earlier study showed that display duration was also not used by females in selecting mates.     

Response to perceived predation threat in fiddler crabs: trust thy neighbor as thyself?

Prey require information if they are to respond to predation threat in a risk-sensitive manner. One way that individuals can obtain this information is through the predator-mediated, threat-induced behavior of conspecifics. We examined such a possibility in a refuge-seeking species, the sand fiddler crab (Uca pugilator). Crabs were either exposed directly to a simulated predation threat (a moving cylinder) or the threat-induced response of a near neighbor. We found that fiddler crabs responded to the flight of their neighbors even when they, themselves, were not privy to the stimulus that induced their neighbors response. However, the wider range of behaviors exhibited by these crabs—which included no reaction, freezing, running back to the burrow entrance, and burrow retreat—suggest that non-threatened crabs either (1) perceived the gravity of the predation threat differently from their directly threatened neighbors and/or (2) engaged in behaviors that allowed them to acquire further information in the face of uncertainty. Conspecific behaviors also had an effect on the hiding duration of crabs, with individuals hiding longer if they saw both the predation threat and the flight of their neighbor. Our results suggest that cues provided by conspecifics can play an important role in guiding the antipredator response of refuge-seeking prey.     

Natal dispersal in two mice species with contrasting social systems

We compared the natal dispersal behaviour of two mice species under laboratory conditions. Natal dispersal is a movement of an animal from its birthplace to its breeding area. This behaviour is known to be influenced by the mating system. In polygamous species, males are more likely to disperse, while in most of the monogamous species, both sexes disperse. Our subjects, the house mouse (Mus musculus) and the mound-building mouse (Mus spicilegus) are two sympatric species of the genus Mus. Both are native in Hungary, but they differ in their habitat type mating system and overwintering strategy. The house mouse is a polygynous species and adapted to human environment, known for mature and reproduce early. On the contrary, the mound-building mice are monogamous, and they inhabit extensively used agricultural fields, where they spend the unfavourable winter period in nest chambers under mounds, which they construct from soil and plant material. Successful overwintering for this species demands delayed maturity and reduced dispersion during the winter. Our results showed that the natal dispersal of these two species differ; both sexes of the mound-building mice dispersed later than the house mice, where a difference between sexes also occurs; house mice males dispersed earlier than females. The mound-building mice showed no sexual dimorphism in this behaviour.     

Spatial and temporal patterns in the feeding behavior of a fiddler crab

Fiddler crabs consume the surficial microphytobenthos around their burrows during low tide. We studied the spatial and temporal feeding patterns in the species Uca uruguayensis by using sequences of digital pictures of feeding pellets accumulation. Data from 61 crabs, feeding without the interference of neighbors, were fitted to different models using Maximum Likelihood and Bayesian approaches. Initial feeding location was independent from the emergence location, and then, crabs continued holding a main feeding direction (clockwise or counterclockwise), suggesting a systematic mechanism that may avoid feeding over already processed sediment. Crabs used at least half of their potential feeding area, but these areas were heterogeneous. Both sexes developed similar feeding areas; however, females were faster and needed less time to feed than males, suggesting that males are time restricted. Our work also highlights the importance of incorporating other underlying mechanisms of the behavior of species into the study of feeding strategies.     

Social influences on territorial signaling in male greater sac-winged bats

Acoustic territorial displays are common among birds but comparatively rare among mammals. An exceptionally vocal mammal well-known for its elaborate territorial displays is the polygynous greater sac-winged bat, Saccopteryx bilineata. Male S. bilineata are often philopatric and establish small territories in their birth colony in which females can roost during the day. During territorial defense, males produce complex territorial songs that are learned through vocal imitation. Territorial songs are mainly produced at dawn and dusk. We studied social influences on male vocal activity and the occurrence of vocal signatures in territorial songs of 27 male S. bilineata from 12 different-sized colonies in Panama. Males produced significantly more territorial songs when they had more territorial neighbors or when they had females roosting in their territories, indicating that male vocal activity rises with increasing male–male competition. Territorial songs are multisyllabic vocalizations with low-frequency buzz syllables being most prominent. We found statistical evidence for a pronounced individual signature encoded in the buzz syllables of territorial songs that could facilitate individual recognition among rival neighbors. Additionally, we found a vocal group signature in territorial songs, suggesting that young males may learn territorial songs from more than one tutor male. Resident male S. bilineata appear to cooperatively defend their colony against male intruders, making a group signature in territorial songs potentially advantageous.     

Density affects female and male mate searching in the fiddler crab, <Emphasis Type="Italic">Uca beebei</Emphasis>

In most species, only one sex searches for mates while the other waits. Models of sex-specific mate-searching behavior predict single-sex searching, but the factors that determine which sex searches are not understood. In this study, we examine the effects of density and predation risk on mate-searching behavior in the fiddler crab Uca beebei. U. beebei is one of the few fiddler-crab species in which both sexes search for mates. In a field experiment conducted in Panama, we manipulated crab density and perceived predation risk in replicate plots. Females searched more and males searched less at high densities. At high levels of perceived predation risk, both sexes similarly reduced their search rates. Observations of plots that naturally varied in crab density show that females were more likely to search for mates in areas of higher density, where there were more males. Females may preferentially search for mates in high-density areas because the abundance of nearby burrows, into which they can run to escape predators, decreases their costs of searching and because the abundance of males and male burrows facilitates comparisons and thus may increase their benefits from searching. Males at high densities decrease their mate-searching rate perhaps in response to the increase in female searching and to the corresponding increase in the intensity of their competitors' mate-attraction signals.     

Sex expression of an immobile coral-inhabiting snail,<Emphasis Type="Italic"> Quoyula monodonta</Emphasis>

To test the hypotheses explaining the sex expression of the immobile snail Quoyula monodonta, which inhabits the surfaces of the branching coral, Pocillopora eydouxi, the size, sex, gonad development, penis length and the composition of neighboring individuals were investigated between November 1994 and August 1995 in southern Taiwan. Although the snails often aggregated and formed patches, more than 50% were solitary. Females were larger than males both within a snail patch and in the whole population, but the overlap in size range was wide. The males were generally accompanied by females, whereas most females were solitary. Females were rarely (6%) found in the same patch with another female, but 35% of the males had male neighbors. Most juveniles found were also solitary. The composition pattern within a patch cannot be explained by random sampling. Gonad development of an individual was dependent on the presence and the sex of its neighbors within the same patch; the penis length of males also depended on the presence of neighbors. These phenomena suggest that an individual is sensitive to its neighbors. No individuals in the process of sex change were ever found from histological studies of the gonads. Neither the hypothesis that the sex of recruits determines their habitats, nor the hypothesis that there is strict protandric sex-change is supported. The results, however, are all clearly compatible with the hypothesis that the snail has labile sex expression. In the presence of existing females in a patch, recruits are more likely to develop into males, whereas recruits starting a new patch grow to a larger size before developing into females directly. The labile sex expression of Q. monodonta is the only such report in neogastropods.     

Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort

We examined how mating success varied in relation to age, weight, body size, and display behavior among great bustard Otis tarda males. The estimated mating success was strongly skewed, with 45% of adult males being involved in copulation attempts and only 9.7% actually seen copulating successfully. Unlike most birds, body size continued increasing in great bustards several years after reaching sexual maturity. Age, weight, and display effort were all significant and independent predictors of male mating success. The higher display effort involved performing longer full-display bouts. Older males could detach from the male flock earlier in the season as well as on each day and spend longer seasonal and daily periods displaying as solitary birds, which contributed to increase their mating success. In contrast, males weighing more did not invest more in display, which suggests that they could be recognized as dominants by other males and selected by females through assessment of their plumage sexual traits. In contrast to most other bird species, the system described for great bustards resembles that found in some lek-mating ungulates, where social rank is a complex trait determined by both age and mass, and as in these mammals, it suggests that sexual selection continues to favor a high male weight in this extremely sexually dimorphic species.     

Sand disposal of the painted ghost crab Ocypode gaudichaudii (Decapoda: Ocypodidae): A possible role in courtship

In 1991/1992, we studied the sand disposal behavior of the painted ghost crabs Ocypode gaudichaudii on the Pacific coast of Panamá. O. gaudichaudii either kick, dump or tamp sand they excavate from their burrows. Here we relate these three kinds of sand disposal to burrow structure and distribution, as well as to crab size and sex. Our objective was to determine whether tamping may be a male courtship signal. Burrows whose owners tamped sand were on average longer, deeper, and higher on the beach than were burrows whose owners kicked or dumped sand. Five burrow shapes were distinguished, with half-spiral and spiral shapes being most common among tamped burrows. All crabs excavated from tamped burrows were males. Tamped burrows peaked in abundance around full and new moons. These observations, together with what is known about mating and breeding behavior of other ghost crabs, suggest that tamping may be involved in O. gaudichaudii courtship.     

A test of the "dear enemy effect" in the strawberry dart-poison frog (<Emphasis Type="Italic">Dendrobates pumilio</Emphasis>)

Some territorial animals exhibit a form of social recognition, commonly termed the "dear enemy effect", in which territory residents display lower levels of aggression toward familiar neighbors compared to unfamiliar individuals who are non-territorial "floaters". Despite the widespread occurrence of territorial social systems and use of acoustic signals for communication in anuran amphibians, only two previous studies have demonstrated vocally mediated dear enemy behavior in a territorial frog. In this study, I conducted neighbor-stranger discrimination playback experiments in a third species of territorial frog, the strawberry dart-poison frog, Dendrobates pumilio (Anura, Dendrobatidae). In the first experiment (n=24), I broadcast the calls of a subject's nearest neighbor and the calls of an unfamiliar individual from the approximate midpoint between the subject's and the neighbor's territories. Although males responded to the stimuli, they did not exhibit differential responses to the calls of neighbors and strangers. In a second experiment (n=22), I broadcast the calls of a neighbor and a stranger to subjects through a speaker located in the approximate center of the neighbor's territory. Males also responded to the playback, although less intensely than in the first experiment, but no discrimination between the calls of neighbors and strangers was found. Thus, territorial males of the strawberry dart-poison frog appear not to discriminate behaviorally between the advertisement calls of neighbors and strangers. Several proximate and ultimate-level hypotheses for this lack of vocally mediated neighbor-stranger discrimination are discussed.Communicated by T. Czeschlik     

Multiple mate choice criteria and the importance of age for male mating success in the microhylid frog, Cophixalus ornatus

We examined multiple mate choice criteria in Cophixalus ornatus, a terrestrial breeding, microhylid frog. Mate choice consisted of three stages: mate attraction (male calling), courtship (male behavior between the call site and the nest), and nest site selection by the female. For male C. ornatus, the possession of a call with low dominant frequency relative to calling neighbors increased the probability that they would attract females. Dominant frequency was negatively correlated with age independent of male mass and snout vent length. When escorting the female from the call site to their nest, males traveled along more convoluted paths than when returning to the nest alone. The convolution of the path was, therefore, considered an aspect of courtship. Females released eggs into nests with structural characteristics typical of nests constructed by older males. Thus, females increased their chances of locating an acceptable nest by preferentially approaching males with lower dominant frequencies. This study is the first to demonstrate that age, independent of mass or snout-vent length, can influence call characteristics in anurans, and it is also the first to demonstrate the importance of male age to female mate choice in an amphibian.     

New birds on the block: new neighbors increase defensive costs for territorial male willow ptarmigan

Previous researchers have hypothesized that site-faithful animals may benefit from the presence of familiar neighbors. This study compares the relative costs of territorial defense against new and former neighbors by male willow ptarmigan (Lagopus lagopus). Territorial defense against new neighbors appeared to require a greater expenditure of both time and effort than did defense against former neighbors. Territorial males that had several new neighbors spent a higher proportion of time fighting than did males with fewer new neighbors, and males with both new and former neighbors spent a greater amount of time fighting with their new neighbors, on average, than with their former neighbors. In addition, fights with new neighbors occurred relatively more frequently and were longer than fights with former neighbors. Finally, fights involving new neighbors tended to escalate to higher levels than fights between former neighbors. Reduced defensive costs for site-faithful, territorial males may provide one explanation for the tendency of males to be more site-faithful than females in many species.     

Parental care and mate attraction in the Florida flagfish, <Emphasis Type="Italic">Jordanella floridae</Emphasis>

Theory suggests that there is a trade-off between mate attraction and investment in parental care. Yet, under some circumstances, such as in species with uniparental male care, it is conceivable that males who provide more care also attract more females, and hence avoid this cost. We tested these two alternative hypotheses using the Florida flagfish, Jordanella floridae, which is a highly sexually dimorphic species with male parental care. We predicted that if components of male care behavior primarily serve offspring survival, they should display these behaviors more in the presence of eggs than without eggs. We also predicted that males should show a trade-off between display, and other activities directed to other fish, and care behaviors. Alternatively, if care has a positive effect on mating success, we expected males to show care even in the absence of eggs, and that they should display more care in the presence of females. We found that care behavior mostly depended on the presence of eggs, males increased care when they guarded eggs, and there was a negative relationship between rates of interaction and care behavior. Most care was exhibited in the absence of other fish while the presence of males and females both had a negative effect on care behavior. There was no difference in care behavior with males or females present. Only nest-visitation rates were highest in the presence of females, suggesting that this behavior is used as part of the courtship. These results support the idea that male care in this species is primarily influenced by natural selection. Males did, however, display fanning and other egg-directed behaviors even in the absence of eggs. The level of these behaviors was constant and independent of display behavior. Hence, it seems that care may be used in mate attraction before the male has received his first clutches.     

The mating system of the brown hyaena,Hyaena brunnea in the southern Kalahari

Summary The mating system of the brown hyaena (Hyaena brunnea) in the southern Kalahari is described. Most brown hyaenas live in small mixed sex groups of related individuals. Some subadults of both sexes leave their natal groups, the females presumably to seek a vacated territory, the males to become nomadic. Mating was only observed between nomadic males and group living females. Group living males display little sexual interest in their own or other group's females, but help to feed the cubs.