Differences in context and function of two distinct waving displays in the fiddler crab, <Emphasis Type="Italic">Uca perplexa</Emphasis> (Decapoda: Ocypodidae)

To respond appropriately to communication signals, animals must have the ability to decipher signal meaning. At a basic level, interpreting the difference between territorial and courtship signals can be vital for the survival and reproduction of social animals. Male and female fiddler crabs communicate with claw-waving displays, but the function of these waves remains uncertain. Species differ in the context in which they wave: Some wave during courtship, some during territorial defence and some during both. In this paper, we provide evidence that males of an Australian species of fiddler crab, Uca perplexa, use two different types of claw waving display, lateral and vertical. Lateral waves are employed solely in a courtship context, whilst vertical waves are employed during courtship as well as territorial interactions. Using video recordings, we show that lateral waves were produced spontaneously (broadcast), and their frequency increased only in the presence of female wanderers. Vertical waves were not broadcast but were elicited by male wanderers during agonistic interactions and female wanderers during close range courtship. Male resident U. perplexa were able to discriminate the sex of wandering crabs on the mudflat at distances of 32 cm. During all resident–wanderer interactions, residents attempted to maintain a position directly between the wanderer and the home burrow and orient themselves to face females and to present the major claw towards males. To our knowledge, this is the first demonstration of the multiple use of waving displays in a fiddler crab species.     

Female territoriality and the function of scent-marking in a monogamous antelope (Oreotragus oreotragus)

In monogamous species, females often choose between males according to the quality of the territories they defend, but the extent to which females themselves contribute to territory defence is frequently underestimated. Here we test for differences in male and female roles during paired scent-marking bouts, a key component of territorial defence, in a monogamous antelope. In two populations (Kenya, Zimbabwe) of klipspringer, Oreotragus oreotragus, both males and females usually scent-marked at the same site, but there were significant differences between sexes in terms of investment within bouts. Females initiated most bouts, thus dictating the marking strategy of the pair. Males initiated relatively few bouts, but deposited more scent marks per bout than females and were usually the last to scent-mark before leaving the site; they marked on the same branches as the female and thus overmarked her scent. Both sexes deposited more marks during paired than solo visits. Immediately preceding and following scent-marking bouts, males approached females and females left males more often than expected. Female scent-marking rates were higher when they were receptive than at other times, and this increase was matched by elevated marking rates of males. Females may increase marking rates when they are receptive in order to test the quality of their mate or to incite male competition. However, these ideas are unlikely to explain female scent-marking behaviour outside the mating season, which appears to be related primarily to territorial defence. We suggest that these differences in investment in scent-marking bouts are consistent with predictions that females may be autonomously territorial and that overmarking of female scent by males is a form of mate-guarding. Received: 17 November 1999 / Received in revised form: 24 February 2000 / Accepted: 13 March 2000     

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.     

Song rates of dark-eyed juncos do not increase when females are fertile

Frequency of singing by birds may vary with reproductive stage in ways that reflect variation in the functions of song in intersexual and intrasexual communication. In dark-eyed juncos (Junco hyemalis) high-amplitude song is produced only by males. To investigate the function of this song, we tested whether fertility of females affected singing by their mates or by neighboring males. Using focal observations, song censuses, and radiotracking data, we determined whether song production varied between and among periods when females were fertile and non-fertile. Our findings show that males do not increase song production when their mates are fertile, nor do they increase song production when neighboring females are fertile. These results suggest that male juncos do not signal their intent to defend territories (or mates) more when females are fertile and that they do not use song to advertise to specific potential participants in extra-pair fertilizations. Received: 13 February 1997 / Accepted after revison: 2 May 1997     

Mating strategies of a nocturnal,desert rodent (Dipodomys spectabilis)

Summary The mating system of a nocturnal, desert rodent, the banner-tailed kangaroo rat (Dipodomys spectabilis) was studied through direct observation, live-trapping, and radiotelemetry over a 13-month period from August 1986 to August 1987. Mating behavior varied from exclusive matings between male and female neighbors to competitive mate searching and direct male competition. In summer matings and early in a November to May breeding season, males located receptive females and mated exclusively with them without disturbance from other males. As the operational sex ratio changed in favor of males, multiple males converged on an estrous female's territory and competed for access to her. However, an older, experienced male usually monopolized the matings of the same one to two close female neighbors for the entire breeding period, and females mated with the same male neighbor over several estrous cycles. Monopolization of females by neighbor males was facilitated by female relaxation of individual territorial defense. Dominant males spent considerable time in the territories of the females they monopolized before and during mating. This relaxation in territorial defense was seen in dyadic encounters in which females tolerated all males but allowed significantly more contact by neighbor than stranger males. Neighbor recognition, therefore, seems important in coordinating the mating interactions of this solitary rodent.     

Non-independent mate choice in a fiddler crab: a case of stimulus enhancement

Non-independent mate selection occurs when the choice behavior of a female is altered by the interactions between other females and males. In the fiddler crab Uca mjoebergi, males court mate-searching females by waving their one greatly enlarged claw. When a female approaches a male, he initiates high-intensity waving. We conducted one natural mate choice experiment and two mate choice experiments using custom-built robotic crabs. We show that the decision of one female to approach a group of males increases the probability that another female will approach and visit a male from the same group. We suggest that this behavior is best explained by the ‘stimulus enhancement’ hypothesis, where the presence of a female near a group of males makes them more likely to be detected by other females due to an increase in male display rate.     

Limitations to colour-based sexual preferences in three-spined sticklebacks (Gasterosteus aculeatus)

Female three-spined sticklebacks (Gasterosteus aculeatus) are suggested to select mates based on their red nuptial coloration, males with a redder display being more preferred. Although there are both laboratory and field data to support this view, there are also published accounts where females do not show a preference for the redder male. Here we report the results of a series of 19 trials where receptive gravid female three-spined sticklebacks were allowed to choose between two size-matched rival males. We used photographic and image analysis techniques to quantify male nuptial coloration to investigate how the magnitude of the colour difference between the two alternative males influenced female preferences. Using the amount of time a female spent oriented towards each male as a measure of his attractiveness to her, females were not always found to select the redder of the two presented males. We did, however, find that that the relative difference in coloration of the two males in each pair was important in determining the level of coloration-based preference, with females only selecting redder males consistently when the difference in coloration was sufficiently large. Received: 26 October 1999 / Received in revised form: 10 January 2000 / Accepted: 13 February 2000     

Determinants of male territorial behavior in a Hungarian collared flycatcher population: plumage traits of residents and challengers

In intrasexual conflicts, contestants can rely on relative or absolute size of status badges to mediate aggressive behavior. Most studies focus on the response of focal animals to variation in status badges of their competitors; few have simultaneously considered the traits of both participants under experimental conditions. By simulating territorial intrusions, we tested the importance of two sexual traits [forehead patch size (FPS) and wing patch size (WPS)] in territorial behavior of males in a Hungarian population of the collared flycatcher, Ficedula albicollis. We presented a stimulus male to an unpaired resident male to examine whether the characteristics of the territory owner or those of the challenger were associated with the latency of the first attack of the owner, which is a good predictor of the territorial behavior in general. WPS of the stimulus male was a significant determinant of the latency of the first attack, as males with a larger WPS elicited quicker attacks from the residents than males with a small WPS. From the residents’ perspective, age appeared to influence their territorial behavior, as yearlings had shorter attack latencies than older males. Additionally, latency could be considered an individual-specific attribute because it varied consistently among males, even when the WPS of the stimulus male was controlled, and it was associated with pairing success. Contrary to findings in a Swedish population, FPS seemed to be unimportant in male–male competition in our population, which suggests population differences in the role of the two plumage traits. Our results indicate that in a territorial conflict, the characteristics of both participants are important.     

Investment in mate choice depends on resource availability in female Galápagos marine iguanas (<Emphasis Type="Italic">Amblyrhynchus cristatus</Emphasis>)

Changes in mate selectivity can significantly alter the direction and strength of sexual selection. When the direct cost of mate search increases selectivity often declines; however, little is known about how the relative cost of mate search affects investment in mate choice. Here, I investigate whether male and female Galápagos marine iguanas (Amblyrhynchus cristatus) alter their investment in mate choice behaviors when resources are limited and the relative cost of mate search is increased. Moderate resource limitation had little effect on male reproductive behavior: in both years, a similar number of males were territorial, and the mean display rate and copulation success of territorial males did not differ. In contrast, female mate search appeared to be affected by the prevailing environmental conditions. During the reproductive season following a moderate El Niño event, when food availability declined, females were in poorer body condition, assessed fewer territorial males, and mated with a male with lower relative reproductive success. Circulating hormone levels also differed between years: when resource availability was limited, receptive females had higher levels of testosterone and stress-induced corticosterone. The frequency and magnitude of climatic fluctuations are expected to increase in the future across many regions of the globe. Determining how sexual selection is shaped by changes in resource availability is vital for predicting the impact of climate change.     

Mating effort and female receptivity: how do male guppies decide when to invest in sex?

Males vary in the degree to which they invest in mating. Several factors can explain this variation, including differences in males’ individual condition and the fact that males allocate their energy depending on the context they face in each mating attempt. Particularly, female quality affects male reproductive success. Here, we studied whether male guppies (Poecilia reticulata) strategically allocated more mating effort, in terms of mating behaviour and male–male competition, when they were matched with a receptive (R) female than a non-receptive one. In accordance with our prediction, we found that males increased their mating behaviour when they were with a receptive female. Even though male guppies can inseminate non-receptive females, we only found high levels of courtship between males that were with a receptive female rather than a non-receptive one. Although there was little affect of female receptivity on male–male competition, we found that males chased and interrupted courtships more with receptive females than with non-receptive females regardless of odour. Finally, we also studied whether the sexual pheromone produced by receptive female guppies is a cue that males use in order to increase their mating effort. We found that males were more attracted to a female when they perceived the sexual pheromone, but only increased their mating and aggressive behaviours when females showed receptive behaviour. This strategic increase in mating effort could result in higher male reproductive success because mating attempts towards receptive females are likely to be less costly and males could have a greater probability of fertilisation.     

Adult female hamsters avoid interspecific mating after exposure to heterospecific males

When females mate with a heterospecific male, they do not usually produce viable offspring. Thus, there is a selective pressure for females to avoid interspecific mating. In many species, females innately avoid heterospecific males; females can also imprint on their parents to avoid later sexual interactions with heterospecific males. However, it was previously unknown whether adult females can learn to discriminate against heterospecific males. We tested the hypothesis that adult females previously unable to avoid interspecific mating learn to avoid such mating after being exposed to heterospecific males. Syrian hamster (Mesocricetus auratus) females not previously exposed to Turkish hamster (Mesocricetus brandti) males can discriminate between odors of conspecific and heterospecific males, but they mate with either type of male. However, when we exposed adult females to both a conspecific male and a heterospecific male through wire-mesh barriers for 8 days, and then paired them sequentially with the two males, females were more receptive to conspecific males and more aggressive to heterospecific males. When females were paired with the heterospecific male first and the conspecific male second, no female was receptive and all were aggressive to heterospecific males. When females were paired with the conspecific male first, only 43% of females were then aggressive toward the heterospecific male. That is, interactions with conspecific males may decrease a female’s ability to properly avoid heterospecific males. Our study clearly shows for the first time that females can learn during adulthood to avoid interspecific mating just by being exposed to stimuli from heterospecific males.     

Sexual behavior is related to badge size in the house sparrow Passer domesticus

Summary Badge size, which functions as a signal of dominance status in male house sparrows Passer domesticus, was significantly related to their sexual behavior. Males with large badges participated in communal displays (multi-male chases directed towards single females) more often than males with small badges, irrespective of whether the female involved was the male's mate or not. Experimentally released females were more often chased if they were fertile than if they were nonfertile. Estradiol-implanted females were chased more often than control females without a hormone implant, and males with large badges chased estradiol-implanted females more often than did males with small badges. Both forced extra-pair copulations during communal displays and unforced extra-pair copulations were more often achieved by males with large than small badges. Male house sparrows with large badges also copulated with their mates at a higher rate than did males with small badges. A higher certainty of paternity therefore is hypothetized to accrue to male house sparrows with large badges.     

Tests of the mate-guarding hypothesis for social monogamy: does population density, sex ratio, or female synchrony affect behavior of male snapping shrimp (Alpheus angulatus)?

There are at least two mechanisms by which social monogamy in the absence of biparental care may evolve: as a form of territorial cooperation, in which one or both sexes benefits by sharing a territory with a partner, and as a form of extended mate guarding, in which males guard females through entire, and perhaps multiple, reproductive cycles. I examined the effects of population variables (density, sex ratio, female synchrony) on male pairing behavior in the snapping shrimp, Alpheus angulatus, to test the hypothesis that social monogamy in this genus has evolved as a result of selection on males for long-term mate guarding of females. There was no evidence that pairing behavior changes with differences in population density; in a natural population, there was a 1:1 relationship between the number in pairs and local population density. In a laboratory experiment, males altered their pairing behavior in response to manipulated differences in sex ratio. Males in female-biased sex ratios were significantly more likely to abandon recently mated females than males in equivalent sex ratios, though there was no significant difference in the duration of pairing or the number of times males switched females. Observations of shrimp maintained for an extended period in the laboratory revealed no evidence that females molt and become sexually receptive synchronously, which would reduce the likelihood that a searching male would encounter additional receptive females. These data suggest that sex ratio may have contributed to the evolution of social monogamy in snapping shrimp, but provide no evidence that population density or female synchronous receptivity have contributed to the evolution of social monogamy.     

Sexual cohabitation as mate-guarding in the leaf-curling spider Phonognatha graeffei Keyserling (Araneoidea, Araneae)

The leaf-curling spider Phonognatha graeffei incorporates a twisted leaf into the central hub of its orb-web that is used as a retreat. This species is unusual among orb-weaving spiders because males cohabit in the leaf retreat with both immature and mature females, mating with the former shortly after the female molts. Cohabitation appears to be a form of mate-guarding because cohabiting males respond agonistically to rival males that venture onto the web, and their behaviour depends upon the reproductive status of the female; males defending immature females are more aggressive than those defending virgin, adult females. Males copulate with previously mated females for significantly longer than with virgin females. Females may cannibalise cohabiting males, which occurs independently of whether the female has been deprived of food. Females that cannibalise a single male do not have a higher fecundity than non-cannibalistic females. Received: 2 February 1996 / Accepted after revision: 27 October 1996     

The mating system of two hybridizing species of water striders (Gerridae)

Summary We studied oviposition, feeding and mating behavior of the water striders Limnoporus dissortis and L. notabilis in sympatric and allopatric populations occurring on semi-permanent ponds in Alberta and British Columbia, Canada. Here we describe the mating system of the species and consider the evolution and maintenance of reproductive strategies employed by both sexes. Females of both species oviposited along edges of floating vegetation, either with a guarding, postcopulatory male in attendance, or alone and independent of copulation. Some females ovipositing alone were discovered by males, and when pressed for copulation, either (1) abandoned oviposition or (2) copulated and then resumed oviposition. Females that oviposited with a guarding male laid more eggs than those that completed oviposition alone. Most breeding females accepted prey offered experimentally, while a large proportion of males rejected prey but responded to model gerrids with aggressive displays and mating behavior. Males displayed three mating tactics: (1) territorial-signaling (TS), (2) patrol-signaling (PS), and (3) silent patrolling (SP). All territorial males produced surface waves from oviposition sites before attempting to mate. Some patrolling males signaled, while approaching potential mates as an aid in sex discrimination, but others did not signal and mounted both males and females. When presented with dead gerrid models, males of L. notabilis discriminated between sexes while those of L. dissortis mounted both male and female conspecifics. Collectively, males employing TS tactics fertilized more eggs than patrolling males and TS males of L. notabilis fertilized relatively more eggs than those of L. dissortis. Individual males switched frequently between territorial and patrolling behavior both under natural conditions and in field exclosures. TS males gave more signals per encounter than PS males suggesting that signaling varies with male dominance. Choice of tactic did not depend upon hunger level and was not associated with significant differences in body length in single-species populations. However, in a mixed population, the smaller males of L. dissortis were rarely territorial and signaled infrequently.     

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     

Male size,mating success,and mating strategy in the mosquitofish Gambusia affinis (Poeciliidae)

Summary In laboratory choice experiments, receptive female western mosquitofish Gambusia affinis affinis (deprived of contact with males 30 days) preferred the larger of two males. When two males differing in size were placed with a receptive female, the larger was generally able to monopolize access to her, but not when the female was not receptive. In other experiments, a single male was placed with a receptive or non-receptive female. These experiments showed size-correlated differences in male mating behavior. Smaller males were more likely to chase non-receptive females and attempt to inseminate them forcibly, while larger males continued to court females even if they were non-receptive. Even when the female was receptive, there was some evidence of size-correlated differences in levels of courtship and chasing.     

Land or lover? Territorial defence and mutual mate guarding in the crimson-breasted shrike

Crimson-breasted shrikes produce duets which are used in interactions with neighbours and intruders. We examined two major hypotheses explaining duetting, the territorial defence and mate guarding hypotheses, using playback experiments. The responses of 12 pairs towards solo male, solo female and stereo duet playback were analysed using principal component analysis and ANOVA. The measures of response intensity revealed in most cases highly coordinated responses of the pair partners. Although these joint responses could suggest cooperative territorial defence or mutual mate defence, responses to the treatments differed between males and females in terms of three variables. Females increased their answer rates significantly in response to unpaired female intruders, compared to unpaired male intruders. During duet playback, females directed their responses equally at both intruding pair members. In contrast, males were equally responsive towards unpaired male intruders and intruding pairs, but in the latter case directed their responses at the male pair member. In the presence of unpaired female intruders, males increased their solo song rates, but in the presence of intruding pairs, they increased overlapping rates and duetted with their mates. These divergent responses suggest that females use duets for mate guarding against unpaired females, whereas males use solo songs for mate guarding and both solos and duets in territorial defence.     

Space use and social structure in meadow voles,Microtus pennsylvanicus

Summary Free-ranging, sexually mature meadow voles (Microtus pennsylvanicus) were tracked by using radiotelemetry from June through August in Front Royal, Virginia, U.S.A. Estimates of intraspecific spacing were derived from the concurrent movements of up to 16 voles. Positions were recorded hourly for 24 h, twice per week. A total of 16 male and 15 female voles were studied during sixteen 24-h sessions.The daily ranges of males (192.3±109.7 m²) were larger and more variable than those of females (68.6±39.4 m²). Males also changed locations more frequently (Fig. 2).Adult females usually maintained territories free of other females; males overlapped considerably among themselves (Fig. 2). Males temporarily moved into the areas occupied by estrous females, indicating intrasexual competition among males for access to receptive females (Fig. 3). M. pennsylvanicus appears to be promiscuous, is socially organized into territorial, maternal-young units during the breeding season, and fits the female territorial model of population regulation.     

A dynamic model of mating behavior in digger wasps: the energetics of male-male competition mimic size-dependent thermal constraints

I develop a state-based dynamic model of behavior to demonstrate that size-dependent differences in temperature tolerances are not necessary to account for the activity of small male digger wasps late in the day. In the model, males defend or patrol the nesting area, wait near nests, or feed away from the nesting area depending on time of day, energy reserves and size rank. I assume a large male competitive advantage, so mating opportunities decrease with size rank for territorial or patrolling males and are rare for all waiting males; the costs of patrolling or defense are higher than the costs of waiting. If energy reserves of all males are initially small, all males alternate feeding and territorial or patrolling behavior. If energy reserves are initially large, large males patrol or maintain territories until they risk starvation and leave the area to feed. At this time, smaller males that have conserved their resources by waiting and feeding may defend territories or patrol. I simulate the behavior of three populations representing two species of Microbembex by assuming large initial energy reserves for populations in which males were territorial and small initial reserves for populations in which males patrolled, and then convert the predicted time of activity to temperature using local regressions from field studies. Temporal patterns in the activity of large and small males were similar to those actually observed, and relationships between size and temperature predicted by the model corresponded to most observations and were sometimes positive. Thus, the delayed activity of smaller males does not correspond to activity at higher temperatures and is probably not attributable to size-dependent thermal tolerances, but may represent a temporal displacement of mating activity due to intra-sexual competition and mediated by energetics. The model makes testable predictions on the timing of feeding and depletion of energy reserves in relation to size and initial energy state, and suggests how differences among species may influence the temporal and spatial organization of male mating behavior. Received: 27 February 1997 / Accepted after revision: 26 July 1997