The dominance of seismic signaling and selection for signal complexity in Schizocosa multimodal courtship displays

Schizocosa wolf spiders show tremendous diversity in courtship complexity, with different species employing varying numbers of components within and across sensory modalities. Using a comparative approach, we investigate the importance of each signaling modality in the courtship display of five Schizocosa species (three stridulating and two drumming) by assessing mating success under manipulated signaling environments. Irrespective of the degree of male ornamentation, the three stridulating species exhibit a dependence on the seismic, but not visual, signaling environment for mating success. Mating was independent of signaling environment for the two drumming species. We next ask whether the degree to which each species depends upon a signaling modality for mating (i.e., modality importance) is correlated with the estimated modality-specific signal complexity. We first calculate effect sizes for the influence of seismic versus visual signaling environments on the likelihood to mate for ten Schizocosa species and then use an element-counting approach to calculate seismic and visual signal complexity scores. We use a phylogenetic regression analysis to test two predictions: (1) the importance of seismic signaling is correlated with seismic signal complexity and (2) the importance of visual signaling is correlated with visual signal complexity. We find a significant relationship between visual signal importance and visual signal complexity, but no relationship between seismic signal importance and seismic signal complexity. Finally, we test the hypothesis that selection acts on complexity per se by determining whether seismic and visual signal complexity is correlated across species. We find support for this hypothesis in a significant relationship between seismic and visual signal complexity.     

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.     

Immune function, dominance and mating success in drumming male wolf spiders Hygrolycosa rubrofasciata

In this study, we provide a piece of experimental evidence that immune function is related to dominance and mating success in wild caught male wolf spiders, Hygrolycosa rubrofasciata. In the mating season, H. rubrofasciata males are actively searching for receptive females, and while searching males often engage in agonistic behavior (i.e., agonistic drumming signals, chases, and fights) with each other. The present results demonstrate that dominant males had higher lytic activities in their hemolymph than subordinates. Lytic activity estimates the concentration of antimicrobial peptides with lysozyme-like activity in hemolymph, which have been shown to play an important role in defense against bacteria, viruses, and fungi. Dominants also had higher courtship drumming rates than subordinates. Moreover, winners in mating competition had higher lytic activities than losers, but this was measured nonindependently of dominance status. Among males with mating failure, there was a moderate negative correlation between encapsulation rate and courtship drumming rate, suggesting that low quality males might not be able to bear the immunological costs of courtship behavior. These results suggest that females might gain immunological benefits by preferring vigorous males.     

Baboon loud calls advertise male quality: acoustic features and their relation to rank,age, and exhaustion

Free-ranging adult male baboons give loud two-syllable wahoo calls during dawn choruses, interactions between groups, when chasing females, and in aggressive interactions with other males. Previous research has shown that the rate and duration of these contest wahoos are correlated with a males competitive ability: high-ranking males call more often, call at faster rates, and call for longer bouts than do low-ranking males. Here we report that acoustic features of wahoos also reveal information about male competitive ability. High-ranking males give wahoos with higher fundamental frequencies (F0) and longer hoo syllables. Within-subject analyses revealed that, as males fall in rank, the hoo syllables tend to shorten within a period of months. As males age and continue to fall in rank, F0 declines, hoo syllables shorten, and formant dispersion decreases. Independent of age and rank, within bouts of calling F0 declines and hoo syllables become shorter. Because wahoos are often given while males are running or leaping through trees, variation in these acoustic features may function as an indicator of a males stamina. The acoustic features of contest wahoos thus potentially allow listeners to assess a males competitive ability.Communicated by C. Nunn     

Female courtship in the Banggai cardinalfish: honest signals of egg maturity and reproductive output?

Despite the vast literature on male courtship behaviour, little is known about the function and information content of female courtship behaviour. Female courtship behaviour may be important in many species, particularly where both sexes invest heavily in the offspring, and if such behaviours contain honest information regarding a females potential reproductive investment, they may be particularly important in male mate choice. Using observations of two female courtship behaviours (the rush and the twitch) from experimental pairings in the Banggai cardinalfish (Pterapogon kauderni), I addressed the question of whether these courtship behaviours contained information on female reproductive output (clutch weight) and egg maturity (proximity to spawning), traits commonly associated with male mate choice. I especially focused on the importance of these courtship behaviours in relation to other female characters, such as size and condition, using multiple regression. I found that one of these behaviours, the rush, was strongly associated with fecundity, whereas size, condition and the twitch were not. Further, I found that the twitch behaviour was associated with how close to actual spawning a female was. The results suggest that female courtship behaviour may convey highly important information in a mate choice context. I discuss the adaptive value of honest information in female courtship behaviour in light of these results.Communicated by K. Lindström     

Nutritional phagostimulants function as male courtship pheromone in the German cockroach, <Emphasis Type="Italic">Blattella germanica</Emphasis>

Summary. Males of the German cockroach, Blattella germanica, secrete a pheromonal substance from the abdominal tergal glands, which elicits a feeding response in females during the sequential courtship behavior. The nuptial secretion consists predominantly of a synergistic mixture of sugars and phospholipids. Cholesterol and a series of amino acids, which are also components of the males glandular secretion, significantly enhanced the phagostimulant activity of the sugar components. The nuptial feeding behavior of the female cockroach is therefore elicited by a complex assortment of nutritive components in the male tergal secretion, including sugars, phospholipids, cholesterol, and amino acids. These results indicate that a mixture of primary metabolites, and not of specific secondary metabolites, serves as a pheromonal cue that appeals to the females gustatory sense and effectively brings her to the precopulatory position. Although the male secretion consists of nutrients, we suggest that these compounds probably do not represent a significant nutrient investment in females and their progeny but rather function as a signal in the mating sequence of B. germanica.     

Bat serenades—complex courtship songs of the sac-winged bat (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>)

Vocalisations of many songbirds, anurans, and insects are shaped by sexual selection. Males acoustically compete for territories, and females choose their mates by means of male courtship songs. In courtship, richness and complexity of elements are often favoured characters. Only a few examples of complex songs are known in mammals. Males of the harem-polygynous sac-winged bat (Saccopteryx bilineata, Emballonuridae) have an uncommonly complex vocal repertoire, and different song types of males are used in the context of territorial defence and in courting females. We classified the daytime vocalisations of 16 male S. bilineata from a colony in Costa Rica, both on the basis of their acoustical properties and the social context in which they occurred. Seven vocalisation types were differentiated: echolocation pulses, barks, chatter, whistles, screeches, territorial songs and courtship songs. Territorial songs were short, rather stereotyped and not obviously directed towards a certain conspecific. They appear to be of importance in male competition for harem territories, in which females roost during the day. Courtship songs were exclusively observed when males displayed towards a female; they were long and complex, and consisted of highly variable elements (calls). We classified the calls in courtship songs of six males into call types, based on acoustical properties, mainly spectral purity and duration. Four call types are described in detail: trills, noise-bursts, short tonal calls, and quasi constant frequency calls. Twelve parameter values were extracted from the most common call type, the trill. Discriminant function analysis of trills showed that different males had different repertoires. This could allow females to use trill parameters for recognition of individual males and thus for mate choice.Communicated by G. Wilkinson     

The respiratory metabolism of Tilapia mossambica (Teleostei)

In a series of experiments 174, 120 and 139 individuals of the teleost Tilapia mossambica (Peters), were acclimated to 30°C and to salinities of 0.4, 12.5 and 30.5, respectively. The effect of temperature and salinity upon oxygen consumption was studied by abruptly transferring fish of different wet weights to temperatures from 15° to 40°C at an average initial pO₂ of 250mm Hg. At each salinity, the proportionate response to temperature is size-independent. The metabolic rate increases as a function of temperature at 15° and 30°C but not at 40°C. Oxygen consumption is, however, salinity dependent; maximum rates are obtained at 12.5S. This salinity is isotonic in the 80 g fish and, to a lesser extent, in the 5 g fish. Reduction in osmotic load is suggested as the probable cause for a greater scope for activity and greater rate of oxygen consumption in 12.5 salinity.     

Copulatory tactics in relation to sperm competition in Rocky Mountain bighorn sheep

Summary Sperm competition arises in the bighorn mating system because receptive ewes are some-times defended and mated in sequence by a series of increasingly dominant rams and because subordinate (coursing) rams commonly succeed in forcing matings with defended ewes. To identify potential components in a male strategy of sperm competition, the social circumstances and schedule of copulation among free-ranging bighorns were investigated. Three such components emerged: (i) Defending and coursing rams copulated with estrous ewes at extraordinarily high rates, presumably to increase the proportion of their own versus rivals' sperm present at fertilization by simple numerical preponderance at insemination. (ii) Coursing but not defending rams concentrated copulations later in the period of estrus (sensu this study). Evidence from domestic sheep suggests that this may correspond to the period of maximum female fertility. A greater risk and/or reduced benefit associated with forced versus cooperative copulation may force coursing rams to restrict copulation to periods of highest fertility, whereas defending rams may gain even from copulations outside this period. (iii) Defending rams copulated at unusually high rates immediately after bouts of successful coursing. Such retaliatory copulations were associated with reduced amounts of courtship and were more likely to occur if the defending ram had copulated infrequently recently. The number of retaliatory copulations was independent of the number of preceding forced copulations, probably because defending rams rationed ejaculates in anticipation of future coursing. The function of retaliation is not clear, although the literature on domestic sheep suggests the possibility of some form of mechanical displacement of rival sperm. Unlike females of many ungulate species, bighorn ewes were typically receptive to cooperative copulation over a period of days. Although persistent courtship by defending rams may coerce such extended receptivity, it is also possible that ewes, facing unavoidable forced matings by coursing rams, gain genotypic benefits for offspring by providing prolonged copulatory access to relatively dominant defending rams.     

Estimating the complexity of bird song by using capture-recapture approaches from community ecology

Repertoire size, the number of unique song or syllable types in the repertoire, is a widely used measure of song complexity in birds, but it is difficult to calculate this exactly in species with large repertoires. A new method of repertoire size estimation applies species richness estimation procedures from community ecology, but such capture-recapture approaches have not been much tested. Here, we establish standardized sampling schemes and estimation procedures using capture-recapture models for syllable repertoires from 18 bird species, and suggest how these may be used to tackle problems of repertoire estimation. Different models, with different assumptions regarding the heterogeneity of the use of syllable types, performed best for different species with different song organizations. For most species, models assuming heterogeneous probability of occurrence of syllables (so-called detection probability) were selected due to the presence of both rare and frequent syllables. Capture-recapture estimates of syllable repertoire size from our small sample did not differ significantly from previous estimates using larger samples of count data. However, the enumeration of syllables in 15 songs yielded significantly lower estimates than previous reports. Hence, heterogeneity in detection probability of syllables should be addressed when estimating repertoire size. This is neglected using simple enumeration procedures, but is taken into account when repertoire size is estimated by appropriate capture-recapture models adjusted for species-specific song organization characteristics. We suggest that such approaches, in combination with standardized sampling, should be applied in species with potentially large repertoire size. On the other hand, in species with small repertoire size and homogenous syllable usage, enumerations may be satisfactory. Although researchers often use repertoire size as a measure of song complexity, listeners to songs are unlikely to count entire repertoires and they may rely on other cues, such as syllable detection probability.Communicated by A. Cockburn     

Mate choice and the supernormality effect in female sticklebacks (Gasterosteus aculeatus)

Summary Female threespine sticklebacks Gasterosteus aculeatus from a marine population were presented simultaneously with two dummy males that differed in size. When the dummies were moved with a carousel apparatus in a way that roughly simulated a male courtship activity (circling) females responded to the dummies with following behaviour normally directed to courting males. Female sexual response was elicited more effectively by the larger (supernomal) dummy, even though it exceeded by 25% the length of the largest males in the study population. Females did not show complete preference for the larger dummy but allocated courtship effort in proportion to the sizes of the two dummies (Fig. 3). The relative projection areas of the two dummies provided the best predictor of how courtship was allocated between them, suggesting a perceptual basis on which females might select mates. This pattern of partial preference for the supernormal dummy occurred in females that exhibited either high or low amounts of following behaviour and is apparently independent of female responsiveness. Supernormality may therefore provide a mechanism whereby secondary sexual traits can be elaborated, even without evolutionary change in preference for those traits by the opposite sex.     

Patterns of the electric organ discharge during courtship and spawning in the mormyrid fish,Pollimyrus isidori

Summary Pollimyrus isidori's electric organ discharge (EOD) is of the pulse type. Patterns of EOD intervals were investigated prior to, during and following spawning behaviors as related with overt behaviors, and with the sound production by the nestbuilding male. Prior to the time of reproduction, isolated and socially interacting fish (n=15) showed characteristic discharge interval patterns for resting, swimming, probing, hovering and hiding activities. Males (n=8) and females (n=6) did not differ in their mean EOD repetition rates during resting (11.6±2.5 Hz), nor Short Bursts/min (less than 20 intervals of 8–13 ms). In interacting fish Long Bursts (greater than 20 intervals of 8–13 ms, lasting for more than 300 ms) were observed only during the attack and bite sequence. A pursuing fish displayed a rapid alternation of Long Bursts with Discharge Breaks (300–1000 ms silence) during the chase behavior. Avoidance behavior which followed from several attacks was correlated with a Medium Uniform Rate (8–12 Hz) normally lasting for 20 to 60 s, or a Discharge Arrest (silence greater than 1 s) in the submissive fish. The nocturnal courtship behavior began soon after dark (1900 h). Spawning typically started 2 to 5 h after dark, continuing for 2 to 6 h until about 0200 h. During courtship and spawning the female's brief visits (15–25 s) to the male's territory recurred every 30–60 s. At all other times the female was aggressively excluded from the nest region. Courtship and spawning behaviors are described along with the electrical displays identified from 19 spawnings in three fish pairs (from a total of 37 spawnings in 4 males and 4 females). Just prior to the onset of courtship behavior, with male territorial aggression beginning to decline, females switched from a Medium Sporadic Rate pattern (resting and hiding patterns; 13 Hz) to a Medium Uniform Rate pattern (6–8 Hz) while still in their hiding area. Females continued to display this uniform rate throughout the courship and spawning period, including the courtship and spawning bouts when Discharge Breaks or Arrests also occurred. This persistance distinguishes the courtship pattern from the similar avoidance pattern (see above). The male courtship and spawning EOD pattern was similar to the female's and unique for a territorial male. He switched from a High Sporadic Rate (swimming EOD pattern; about 18 Hz) to a regularized Medium Uniform Rate (about 9 to 11 Hz) only during courtship and spawning bouts, including 1–3 EOD Breaks during Vent-to-Vent coupling (average interval: 272±71 ms, n=37). No sooner had the female left the spawning site than he resumed displaying a High Sporadic Rate. This temporal correlation of reproductive behaviors with electrical displays suggests their instrumental role in mutual acceptance of mates. Males showed their sex-specific type of EOD phase-locking, the Preferred Latency Response, only during the first few hours of entry of a fish in their tank. Two females with EOD waveform features more typical of males also spawned repeatedly; waveform does not appear to be critical. Males stopped their nocturnal sound production for the later part of courtship and the whole spawning period. Except for infrequent attacks on the female between spawning bouts, the male did not resume singing until the end of spawning when all eggs were shed (around 0200 h); from this time on the male sang until dawn. The sequencing of the three acoustic elements (moans, grunts, growls) are described. A catalogue of discharge patterns correlated with overt behaviors (Tables 1, 2), and an integrated summary time table of P. isidori's complex reproductive behavior are presented.     

Active mate choice at cock-of-the-rock leks: tactics of sampling and comparison

Summary The importance of active mate choice as a component of sexual selection remains controversial, despite recent studies identifying traits that correlate with male mating success. We analyzed the sampling sequences of individual female Guianan cock-of-the-rock, providing quantitative documentation of active female choice in this lekbreeding species. We compared observed sampling patterns with null models of random female movement and of movement weighted by the observed distribution of male success. The courts actually visited by individual females were significantly more clumped and centrally located than predicted by computer simulations of either random or weighted movement. Contrary to simplified hypotheses of female choice, hen cock-of-the-rock were not limited either to sequential comparisons or to threshold decision rules when selecting mates. Instead, they repeatedly sampled a subset of potential mates, using the pool-comparison tactic. Within this pool, females preferentially mated with the males ranked highest in terms of total courtship visits received, and rejected low-ranking males. Our results suggest complex interactions between patterns of female sampling and male dispersion in lek mating systems, and also provide insights into the constraints on female sampling in lek species.     

Agonistic interactions between male Frontinella pyramitela (Araneae,Linyphiidae)

Summary Adult male bowl and doily spiders, Frontinella pyramitela (Walckenaer) (Linyphiidae), occupy females' webs during courtship, mating, and foraging. When males are abundant, they can be found on about 20% of all adult females' webs, and encounters between the territorial resident males and intruder males occur frequently. Stereotyped interactions take place in 78% of encounters when the resident male has recently copulated on the web, though the presence of the female during the encounters has no effect. When the resident male has had no recent copulatory experience on the web, only 33% of encounters evoke visible stereotyped interaction.Analyses of videotaped interactions reveal that transitions from one behavior to another, though non-random, are highly variable (Fig. 1). The overall ordering of the interactions is species-typical with respect to the relative frequency of behavioral units and the relative amount of time spent in each one.Outcomes of the stereotyped interactions are very predictable: the larger male wins in 75% of all interactions, though when male sizes are closely matched, some significant resident advantage is detectable. A weak but significant inverse correlation between mass difference and duration of the jawlock display (Fig. 2) suggests that the display provides each male with information about his opponent's relative size.     

Female response to song reflects male developmental history in swamp sparrows

One explanation for why female songbirds attend to male song is that the quality of a male's song is associated with the quality of his developmental history. We tested this hypothesis by playing back to female swamp sparrows (Melospiza georgiana) songs recorded from males of either inferior or superior developmental histories, as assessed by their rates of mass gain during the first 18 days post-hatching. Females showed significantly higher levels of courtship display in response to songs of males with superior growth than to songs of males with inferior growth. Out of nine song traits measured, only song duration correlated with variation in female response; duration was also the only trait that differed significantly in univariate comparisons between the superior growth songs and the inferior growth songs. In a multivariate analysis, however, inferior growth songs were best discriminated from superior growth songs by combining three song traits: trill rate, stereotypy, and the number of notes per syllable. We suggest that early developmental stress degrades song in many small ways, and that it is the cumulative effect of the resulting deficits that explains lower female response.     

Courtship and reproduction in<Emphasis Type="Italic"> Carybdea sivickisi</Emphasis> (Cnidaria: Cubozoa)

Courtship and fertilization events in cubozoans have received little attention from biologists, and much of what we know about these processes is based on conjecture or scant anecdotal evidence. I set out to describe these processes in the cubozoan Carybdea sivickisi by observing mature medusae in vitro. Mature adults engage in courtship during which spermatophores are transferred from the male to the female, who then inserts the gametes into her manubrium. Females accepted multiple spermatophores from multiple males, and only produced one embryo strand. This study also provides evidence that the presence of conspicuous velar spots on the females bell margin is a signal of sexual maturity, and that sexual maturity was not reached in either sex until individuals had a bell diameter of at least 5 mm.Communicated by R.J. Thompson, St. Johns     

Gene flow among giant clam (Tridacna gigas) populations in Pacific does not parallel ocean circulation

Tridacna gigas (L.) populations were sampled from 13 locations throughout the west Pacific in 1990–1991 and screened for allozyme variation at seven polymorphic loci. Significant genetic differences were found between the few large stocks of T. gigas remaining in the world. Cluster analysis identified two major regional groups of populations: a Central Pacific group consisting of Kiribati and the Marshall Islands, and a West Pacific group comprising all other populations. Within the latter group, the Great Barrier Reef populations were significantly differentiated from those in the Solomon Islands. The main routes of gene flow did not run parallel with known major surface currents as might be expected, but crossed them. Gene exchange was limited east-west between the Central and West Pacific groups and the GBR and the Solomon Islands. The lack of correspondence between the major surface currents and the routes of gene flow suggests that the genetic structure of T. gigas reflects historical patterns of migration that no longer occur, rather than presentday dispersal. These findings raise fundamental questions concerning the origin and maintenance of genetic diversity in Pacific faunas, and have important implications for the mariculture and conservation of T. gigas.     

Sexual signalling and viability in a wolf spider (Hygrolycosa rubrofasciata): measurements under laboratory and field conditions

This study examined the crucial prediction of the conditional-handicap theory, the relationship between male sexual trait size and male viability, in the wolf spider Hygrolycosa rubrofasciata. In this species, males court females by drumming dry leaves with their abdomen, and males with the highest drumming rate enjoy highest mating success. We determined male drumming rate, body mass, and mobility, which reflects mate-searching activity, in relation to male survival. Because it is often difficult to know how results obtained from laboratory studies reflect the natural world, particularly when the measured variable is survival, we repeated our study in both laboratory and field conditions. Males drumming at the highest rate survived better than males drumming at a lower rate in both laboratory and field conditions. These results are in accordance with the predictions of conditional-handicap models of sexual selection. Survival was independent of male body mass and there was no significant correlation between male drumming activity and body mass. However, large males moved further than smaller males, and males moving longer distances lost less mass than those moving shorter distances. These results suggest that, moving, and consequently mate-searching activity, may be a condition-dependent trait and that there may be a advantage for large males in mate searching. Received: 22 October 1998 / Received in revised form: 28 January 1999 / Accepted: 14 February 1999     

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.