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     

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.     

Temporal constraints and female preference for burrow width in the fiddler crab, <Emphasis Type="Italic">Uca mjoebergi</Emphasis>

We studied sampling behaviour and mate choice in the fiddler crab Uca mjoebergi. Once a female selects a mate, she copulates in his burrow and remains there until releasing her aquatic larvae. U. mjoebergi occurs in habitats that are inundated only by the highest amplitude spring tides. Females can only release their larvae during these tides, and release before or after will result in complete failure of reproductive effort. Matings occur over a 5-day period near the end of neap tides. Our results suggest that within the mating period, females adjust their larval developmental rates by selecting specific burrows in which to incubate their clutches. We found that at the start of the mating period, females chose larger males as mates. Since male size was positively correlated to burrow width, females were selecting wide burrows and effectively incubating at lower temperatures. This would slow down the developmental rates of larvae. In contrast, females that mated late in the mating period selectively chose small males. By incubating in narrower, warmer burrows, these females may increase the developmental rates of larvae. We propose that females are selecting burrows to influence incubation rate and ensure timely release of their larvae. Female U. mjoebergi appear to adjust their preference for the direct benefits of mate choice to increase their reproductive success.     

Experience with chemotactile cues indicating female feeding history impacts male courtship investment in the wolf spider <Emphasis Type="BoldItalic">Schizocosa ocreata</Emphasis>

Male fitness is often determined by the ability of the male to gain access to multiple mates, although in species that exhibit sexual cannibalism, males might increase their likelihood of being cannibalized with each encounter. This risk should create selection for males who are able to perceive potential risks associated with mating encounters. We studied male Schizocosa ocreata wolf spiders to determine whether they use female chemotactile cues (silk, excreta) as indicators of potential risks and how these cues affected subsequent male courtship behaviors. Female treatments included satiated females vs. starved females, as well as a treatment where females had recently cannibalized a male S. ocreata. We performed experiments to assess (1) if males use female chemotactile cues to determine potential risks associated with differing female feeding treatments and alter courtship investment, and 2) how male experience with female chemotactile cues affected courtship investment in subsequent female encounters. At first encounter, males do not vary courtship investment (number of bouts, duration, and vigor) with different female feeding treatments. However, male behaviors during subsequent encounters with female chemotactile cues varied in complex ways, depending on female feeding treatment and male first encounter experience. These data suggest that male experience impacts perception of female chemical cues and offers the male opportunity to avoid sexual cannibalism in high-risk situations.     

Male remating and female fitness in the wolf spider Pardosa astrigera: the role of male mating history

Although the effects of male mating history on female reproductive output and longevity have been studied in insects, few such studies have been carried out in spiders. In a mating system in which females are monandrous while males are polygynous, females may incur the risk by mating with successful males that have experienced consecutive matings and suffer from the possible depletion of sperm and/or associated ejaculates. Here, we examine the effects of male mating history on male courtship and copulation duration, female reproductive fitness, and female adult longevity of the wolf spider, Pardosa astrigera. Results indicated that male mating frequency had little effect on their subsequent copulation success, and of 35 males tested, about half of the males were able to copulate with five virgin females successively at an interval of 24 h. Male mating history had little effect on their courtship duration. However, male mating history significantly affected male copulation duration, female adult longevity, and reproductive output. Males that mated more frequently copulated longer and more likely failed to cause their mates to produce a clutch, although there was no significant difference in the number of eggs laid and the number of eggs hatched regardless of the first clutch or the second one. Multiple mating of male P. astrigera resulted in significant reduction in female adult longevity. Our results indicate that monandrous females mating with multiple-mated males may incur substantial fitness costs.     

Effect of visual background complexity and light level on the detection of visual signals of male <Emphasis Type="Italic">Schizocosa ocreata</Emphasis> wolf spiders by female conspecifics

For visual signaling to be effective, animal signals must be detected and discriminated by receivers, often against complex visual backgrounds with varying light levels. Accordingly, in many species, conspicuous visual displays and ornaments have evolved as a means to enhance background contrast and thereby increase the detection and discrimination of male courtship signals by females. Using video playbacks, we tested the hypothesis that visual courtship displays and leg decorations of male Schizocosa ocreata wolf spiders are more conspicuous against complex leaf-litter backgrounds. Video exemplars of courting males with manipulated leg tufts were superimposed on different backgrounds (complex leaf litter in sun or shade, featureless gray background) and presented to female spiders. Females were more likely to orient to males presented against lighter backgrounds (litter in sun, gray) than the darker ones (litter—shade). Males with larger tufts were also more likely to be detected, as latency to orient was shortest for enlarged and longest for removed tufts. Latency of females to approach was shorter against lighter backgrounds, and approach latency was longest for males without tufts. Female receptivity scores were significantly greater for males against lighter backgrounds, and males with larger tufts had higher scores. These results suggest that both complexity and light level of display backgrounds affect the detection of male visual courtship signals by females and that aspects of the male phenotype may increase chances of detection (and receptivity) against visually complex backgrounds.     

Species recognition and species isolation in wandering spiders (Cupiennius spp.; Ctenidae)

Summary One of the roles ascribed to spider courtship is that of mate recognition and reproductive isolation. We describe behavioral mechanisms underlying these functions in three species of wandering spiders of the genus Cupiennius (Ctenidae). C. salei and the two sympatric species C. getazi and C. coccineus discriminate conspecifics and heterospecifics in a multistage process which covers three principal phases of courtship: (1) Chemical phase: male spiders initiate vibratory courtship communication less often upon contact with heterospecific than with conspecific female silk. (2) Vibratory phase: females respond with their own vibration less often and with increased delay to the vibratory courtship signals of heterospecific males than to those of conspecific males. (3) Tactile phase: in heterospecific pairings males and females interrupt the contact phase if their courtship has progressed thus far. The product of the probabilities of passing through each of the different stages of courtship is between 0.44 and 0.88 for conspecific pairings, but zero for most heterospecific pairings. This is to a greater extent due to the behavioral barriers of the vibratory and the tactile phase than to that of the chemical phase. Male taxis on tracks of female silk does not depend on the species-specificity of the silk. Female vibrations are not essential for species recognition: males increase their signal rates, orient toward and approach responding females in both con- and heterospecific pairings. Female attacks on males are similar to attacks on prey; males defend themselves with outstretched front legs and are caught and killed by the female in only 5% of the cases. In all phases of courtship, female C. salei are least and female C. getazi most selective. When female C. coccineus and C. getazi are involved in heterospecific pairings no copulation is attempted. In contrast, 13 copulations of C. salei with males of the other two species were observed; egg sacs never resulted from these copulations. Among males, those of C. getazi were least selective. The likelihood of heterospecific pairs passing through all phases of courtship is higher in case of the allopatric than in that of the sympatric species. This finding applies to the courtship as a whole, but not necessarily to each of its phases.Offprint requests to: F.G. Barth     

Assessments and decisions in Metellina segmentata (Araneae: Metidae): evidence of a pheromone involved in mate guarding

Assessments and decision-making underlying the initiation of mate guarding in a common web-building spider, Metellina segmentata, are examined in a series of field and laboratory studies. Adult males do not build webs but wander in search of females and mating opportunities. Adult males then wait at the edge of the webs of females and guard them prior to courtship and mating. Guarded females were heavier, larger and carried more mature eggs than solitary females. An active process of information gathering is apparent from introductions of males to the webs of females. Males make accurate assessments about female quality, even in the absence of the resident female. Cues involving web architecture are not used. Males may assess pheromonal cues on the web of the female in deciding whether to guard or abandon a female.     

Limb regeneration and subsequent asymmetry in a male secondary sexual character influences sexual selection in wolf spiders

Males of the brush-legged wolf spider Schizocosa ocreata (Hentz) have conspicuously decorated forelegs used in courtship and agonistic displays. Approximately one in five juvenile males has a missing or regenerating foreleg, and regeneration of a leg lost during development usually results in the absence of a decorative tuft on that leg. The subsequent asymmetry in this male secondary character significantly decreases success in both courtship of females and male-male agonistic interactions. Experimental removal of tufts from one leg of previously successful symmetric males produces similar results. As a test for concomitant behavioral effects, female spiders were shown video images of a courting male with symmetric tufts and the same video image altered to have asymmetric tufts. Female receptivity to the asymmetric video image was lower. In contrast to fluctuating asymmetry resulting from developmental instability, leg tuft asymmetry in S. ocreata most likely arises from a single event during ontogeny – possibly leg loss from an aggressive or predator encounter  – and may serve as a quality indicator in female mate choice. Received: 27 July 1995/ Accepted after revision: 19 November 1995     

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     

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     

Role-reversal in katydids: Habitat influences reproductive behaviour (Orthoptera: Tettigoniidae,Metaballus sp.)

Summary Mating behaviour of the katydid Metaballus sp. varies. At sites QS and DB (in 1982) females competed for access to calling males and males chose mates by mating with heavier, more fecund females. At another site (BR) there was no evidence of role-reversal in reproductive behaviour, and males were observed to compete for mates. This species has a large spermatophore, a product of male reproductive glands, that is eaten by the female after mating. Males at the DB site had small reproductive glands. This suggests that some aspect of the QS and DB environments decreases spermatophore production; spermatophores become a limiting resource for females resulting in the reversal in reproductive roles observed at these sites. A field experiment that involved moving individuals from site BR to QS in 1983 determined that mating system was influerced by site (Table 1). At BR, males produced a continuous calling song, a third of the males observed attracted mates, and called for about 30 min before the female arrived; courtship duration was short. Males that were moved from BR to QS encountered a higher density of receptive females as all males attracted females after an average of just 3 min of calling. They changed their behaviour by producing short periodic bursts of song (zipping), and by courting females for long periods of time. The long courtship period may function as as a mate-assessment period for males. The reproductive behaviour of BR males moved to QS differed from that of native QS males only in the length of time spent in copula.     

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.     

Sexual selection,male morphology,and the efficacy of courtship signalling in two wolf spiders (Araneae: Lycosidae)

Males of the brush-legged wolf spider, Schizocosa ocreata (Araneae: Lycosidae), possess a conspicuous male secondary sexual character: dark pigmentation and tufts of bristles on the tibiae of their forelegs. We tested several hypotheses relating to the role of this conspicuous trait in sexual selection. Triad mating experiments suggest that the tufts do not play an obvious role in the operation of sexual selection by either male competition or female choice, as there were no significant differences in the mating success of intact and experimentally shaved males. However, females mated more often with males that initiated courtship first, suggesting that capture of a female’s attention by male signalling may play a critical role. In behavioral experiments that paired a single male with a female in arenas that allowed both visual and vibratory signal transmission during courtship, female receptivity did not vary significantly with the presence or absence of tufts. However, experiments that isolated the visual component of communication (by eliminating vibratory communication) revealed a significant effect of the presence of tufts: females showed receptivity less often to males with tufts removed. Female response to visual signals was much greater in S. ocreata than in its sibling congener, Schizocosa rovneri, which lacks male tufts. We hypothesize that the tufts serve to increase the efficacy of visual displays of S. ocreata, as vibratory communication is constrained by the complex leaf litter habitat of some populations. Such environmental constraints may make visual signalling over distance a critical factor for effective courtship communication, which may in turn strongly influence male fitness. Received: 30 September 1994/Accepted after revision: 4 August 1995     

Parental investment, potential reproductive rates, and mating system in the strawberry dart-poison frog, Dendrobates pumilio

We studied the effect of relative parental investment on potential reproductive rates (PRRs) to explain sex differences in selectivity and competition in the dart-poison frog Dendrobates pumilio. We recorded the reproductive behavior of this species in a Costa Rican lowland rainforest for almost 6 months. Females spent more time on parental care than males, and `time out' estimates suggest that PRRs of males are much higher than than those of females, rendering females the limiting sex in the mating process. Males defended territories that provide suitable calling sites, space for courtship and oviposition, and prevent interference by competitors. Male mating success was highly variable, from 0 to 12 matings, and was significantly correlated with calling activity and average perch height, but was independent of body size and weight. Estimates of opportunity for sexual selection and variation in male mating success are given. The mating system is polygamous: males and females mated several times with different mates. Females were more selective than males and may sample males between matings. The discrepancy in PRRs between the sexes due to differences in parental investment and the prolonged breeding season is sufficient to explain the observed mating pattern i.e., selective females, high variance in male mating success, and the considerable opportunity for sexual selection. Received: 9 June 1998 / Received in revised form: 27 March 1999 / Accepted: 3 April 1999     

Female sex pheromone in immature insect males—a case of pre-emergence chemical mimicry?

Sexual selection by competition for mates is a formidable force that has led to extraordinary adaptations in males. Here we present results suggesting a novel case of pheromone mimicry in males of Lariophagus distinguendus, a parasitic wasp of beetle larvae that develop in stored grain. Females of L. distinguendus produce a pheromone even before they emerge from a grain. Males are attracted to the parasitised grain and wait for females to emerge. Males emerging later than others are under enormous selection pressure since females mate only once. We show evidence that developing males fool their earlier emerging competitors by mimicking the female pheromone. Males exposed to pupae of either sex exhibit typical courtship behaviour. Searching males are not only arrested by grains containing developing females but spend as much time on grains containing developing males. Hence, by distracting their competitors away from receptive females late males may increase their own chance to mate with these females. After emergence, males decompose the active compounds within 32 h probably to decrease molestation during their own search for mates. Chemical analyses of active pheromone extracts and bioassays using fractions demonstrate that the active compounds are among the cuticular hydrocarbons.     

How do weaponless male fiddler crabs avoid aggression?

Mimicry of females enables weaker males in many species to avoid intrasexual aggression. In fiddler crabs (Uca annulipes), males use their major claw in aggressive interactions to acquire and defend a territory. Males that have autotomised their major claw will be disadvantaged in fighting, but might use their temporary resemblance to females to avoid costly aggressive encounters with other males. We investigated whether: (1) courting males mistake clawless male fiddler crabs for females; (2) clawless males are able to acquire, defend and retain territories as successfully as intact males; and (3) clawless males are more cautious than intact males. Clawless and intact males differed in burrow acquisition methods and fighting behaviour, but were equally successful at acquiring and retaining burrows. While courting males treated clawless males as female, we found no evidence that clawless males mimic the behaviour of females, or that they exploit the advantage of their mistaken identity. Clawless males further appear to avoid male aggression by altering their territorial strategies to minimise the potential for conflict.     

Male mating preference is associated with risk of pre-copulatory cannibalism in a socially polymorphic spider

We performed male attraction experiments and staged courtship sequences to test for non-random mating with respect to social behavioral phenotype in the comb-footed spider, Anelosimus studiosus. While asocial behavior is the dominant phenotype in all A. studiosus populations examined to date, a social phenotype approaches a frequency of 15% in colder environments. We collected test subjects from higher latitude polymorphic populations and scored all individuals as to their behavioral phenotype prior to their use in these trials. Males of both phenotypes differentially approached and courted social females over asocial females and no-spider controls. By offering males different numbers of females of one type vs. the other in subsequent trials, we determined that the difference in attractiveness between the two phenotypes social/asocial is 1.5/1. Both the web produced by a female and a female that has been removed from its web attract males. We suggest that the male attracting pheromone is present on females and is also attached to silk threads. Staged encounters completed between males and females of the respective phenotypes demonstrated that courting males suffer significantly less pre-copulatory sexual cannibalism with social females than with asocial ones, and thus, female social tendency is phenotypically linked to sexual aggression. We propose that the male preference for social females is adaptive because of the observed asymmetry in courtship success.     

Mating system of the burrowing crab Neohelice granulata (Brachyura: Varunidae) in two contrasting environments: effect of burrow architecture

Few studies conducted in crustaceans have demonstrated how habitat features could shape the mating systems. Here, the burrow of Neohelice granulata was considered as a resource used for mating, and its architecture was characterized in two contrasting study sites: Mar Chiquita Lagoon (MCL), an estuary composed of muddy sediments, and San Antonio Oeste (SAO), a marine bay composed of gravel sediment. Burrow features differed between study sites and occupant gender. Large males constructed burrows with a chamber in MCL and with a widened entrance in SAO, while small males constructed the same narrow burrows as females at both study sites. Field experiments demonstrated that burrows with chambers or widened entries are places used for copulation, although successful post-copulatory guarding was displayed only in those with chambers. The intensity of the agonistic encounters and the success of males in winning resources (burrows/females) also depend on the habitat characteristics. N. granulata shows a resource defense mating system where males employ different mating strategies according to the burrow architecture to ensure mating success.     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000