Female-enhanced male competition determines the first mate and principal sire in the spider Linyphia litigiosa (Linyphiidae)

Summary Two to five days before sexual maturation, female sierra dome spiders (Linyphia litigiosa: Linyphiidae) undergo a transformation in their behavior toward males that visit their webs. During this latter part of their penultimate instar, females change from consistently positioning themselves far away from males to actively maintaining close proximity, reactions I call avoidant and associative behavior, respectively. Consistent associative behavior ceases after the female's first mating and thus is limited to soon-to-mature penultimate females. When a mate-seeking male fords an associative female, he attempts to guard her until she matures; this is often a multi-day affair. In contrast, males guard immature avoidant and mature mated females for only a single day. This dichotomy in male guarding times can be understood by the fact that associative behavior signals that the female will soon develop peak reproductive value. Upon completion of their final molt, 98% of females immediately mate with the current guarding male. Secondary suitors are not as likely to achieve mating. Moreover, first mates father 1.8 times more offspring, on average, than secondary mates. Whenever they meet on any female's web, males fight until one of the contestants withdraws. Fights typically are intensive, occasionally deadly, and often result in usurpation of the web by the newly arriving male. Larger males win more fights, but other qualities (e.g., vigor and persistence) appear to be important when contestants differ by less than 10–20% in body weight. Prolonged (i.e., multi-day) guarding of associative females enhances the intrasexual selection process by ensuring that every male that arrives at the web finds it already guarded. Therefore every male that finds the web becomes a participant in a series of male-male conflicts and web usurpations which span the period between the resident female's commencement of associative behavior and her sexual maturation. Since unforced male departures from the webs of associative females are rare, victors are retained on the web until they themselves lose a fight. This facilitates a steady increase in the fighting ability of sequential guards throughout the associative period, up until female maturation and mating. On my study site, first mates represented the final winners in a combative sorting process based on a minimum average of 2 fights; they were heavier and larger than secondary mates and randomly sampled males. The combination of (1) associative behavior by nearly mature females, (2) high mating propensity of newly mature females, and (3) first male sperm priority, constitutes a system whereby females enhance male-male competition and boost the expected fighting prowess of the principal sire of their progeny. Since males appear to make no material contribution toward progeny, the female's behavior probably functions to improve the genetic constitution of the offspring. In addition, the timing of associative behavior may limit prolonged guarding by food-stealing males to a period (1) encompassing the female's pre-molt fast and (2) before the heavy yolking of eggs, thereby ameliorating the nutritional costs of intrasexual selection.     

Idiothetic orientation of a wandering spider: Compensation of detours and estimates of goal distance

Summary The wandering spider Cupiennius salei Keys uses idiothetic orientation, i.e., memorized information about its own previous movements, to retrieve lost prey. Spiders, having been chased away from a prey fly, return to the capture site (the goal) over a distance of more than 75 cm even though all external orientation cues were precluded. This behavior and its sensory basis were examined by varying the proprioceptive and motor command inputs to the memory and by ablating particular lyriform slit sense organs on the legs of the spider.The success rate of returns to the goal after rectilinear chases over 6 discrete distances ranging from 20 cm to>41 cm declines with increasing distances. At distances>41 cm, more than 50% of the performances of intact spiders are nevertheless successful, in that the animals approach the capture site as close as 5 cm (or less).Animals that have been operated on (lyriform organs on all femora destroyed) are much less successful even at short distances. The mean starting angles of the returns by intact spiders and by those operated on do not differ signficantly. Walking error e _n for each segment of the entire return path shows that intact animals deviate little from the ideal return route and correctly estimate the distance to the goal. The operated spiders tend to drift off the ideal return route, while their distance estimates remain largely accurate.Returns after curvilinear chases through a semicircular corridor do not retrace the curved path; instead the spiders take a shortcut. Of all performances by intact and by control spiders (with sham operations) 85% are successful. By contrast, most of the 8 groups with sensory ablations have a success rate of less than 50%.Compensation for the semicircular detours is not quite complete: the mean starting directions of returns are biased, pointing to the corridor, and the shape of many return paths reflects the curved corridor shape. Spiders with unilateral ablations of their femoral lyriform organs show low success rates only if the operated legs are on the inner curve perimeter during the chase, while their return parameters resemble those of the intact group in the reverse situation (operated legs on outer perimeter). These side-specific ablation effects, which are correlated with the geometrical situation existing while idiothetic information is gathered and memorized, suggest that the idiothetic memory depends at least partly on input from proprioceptors.     

Solar orientation in the snail Nerita plicata (Prosobranchia: Neritacea) on a beach near Watamu,Kenya

Investigations carried out under natural conditions suggest that Nerita plicata Linné is capable of orientating to the sun. Behaviour is predominatly photonegative during the morning, and photopositive during the afternoon. Accuracy of orientation improves as the brightness of the sun increases. At Watamu (Kenya), snails tended to orientate away from the sea at all times of the day. Orientatory performance is not affected by the temporal proximity of high or low water, and locomotory speeds seem to be unaffected by the strength of the sun, the time of day, or the state of the tide.     

Biparental defence in a spider mite (Acari: Tetranychidae) infesting Sasa bamboo

Summary Field observations on the nesting pattern of Schizotetranychus celarius (Banks) (Acari: Tetranychidae) revealed that many individuals live gregariously in a united nest. Behavioural experiments recorded on video tape recorder and 16 mm film showed that adult males and females can effectively defend their nest and offspring from a phytoseiid predator, larvae of Typhlodromus bambusae Ehara (Acari: Phytoseiidae), through counterattacks. The male spider mite often killed the predator's larva and succeeded in defending offspring and nest completely. Comparison of the behaviour patterns recorded between a nest owner parent S. celarius and two kinds of intruders (predators and conspecific males or females) revealed conspicuous differences. I suggest that there is a kind of biparental care in S. celarius, and that the life-pattern of this species therefore fits the definition of subsociality originally framed for insects.     

Solar and landscape cues as orientation mechanisms in the beach-dwelling beetle Eurynebria complanata (Coleoptera,Carabidae)

The present paper reports on the first orientation experiments conducted on the strand-living beetle Eurynebria complanata (Linnaeus, 1767) (Coleoptera, Carabidae). The experiments were carried out from June to October 1988. Two different populations were used: one inhabiting a Tyrrhenian beach in Italy and the other a beach on the Atlantic coast of France. Response to sun compass cues was demonstrated in each population at the collection site and for the French population after transportation to Italy, where experiments were carried out on a differently oriented beach. Landscape cues were shown to improve the beetle's orientation capacity when these were tested together with a visible sun. Artificial landscapes were also tested under laboratory conditions. Different-sized silhouettes were placed in the four cardinal directions, and these envoked different responses depending on their height. The beetles oriented towards a black silhouette when this subtended an angle of 25°. The results obtained for the two populations are compared and discussed from an eco-ethological point of view.     

Male signaling behavior and sexual selection in a wolf spider (Araneae: Lycosidae): a test for dual functions

Male signaling behaviors are often studied in a single context but may serve multiple functions (e.g., in male–male competition and female mate choice). We examined the issue of dual function male signals in a wolf spider species Schizocosa ocreata (Hentz) that displays the same species-specific signaling behaviors in both male–male and male–female contexts. These signaling behaviors have been described as either aggression or courtship according to the context observed. We tested the possibility of dual functions by comparing the relationship between behaviors and outcome of male–male contests (winner/loser) and male–female mating encounters (mating success). Frequency, rate, and mean duration of signaling behaviors did not vary with outcome of male–male contests, which appears instead to be based upon relative size and body mass. Winners of contests had significantly greater body mass than losers, and greater mass relative to opponents was significantly associated with probability of winning. Overall, signaling rates were much higher in male–female interactions than in male–male contests and were higher for males that successfully mated than for those that did not mate. Mean duration of some male displays was also greater for males that successfully mated. However, male size was not associated with probability of mating. Taken together, results suggest an intersexual selection context for the current function of male signals in these wolf spiders and that increased display vigor is associated with male mating success.     

Fissioning minimizes ranging costs in spider monkeys: a multiple-level approach

The adjustment to deal with intragroup food competition is probably the most plausible explanation of high levels of fission–fusion dynamics. However, studies did not always support expected relations between food availability, ranging costs, and subgroup size. We used several levels of analysis differing in the time and spatial scale in order to investigate this explanation in spider monkeys. In our study, subgroups were larger when food availability was higher across most levels of analyses used. We also found a fine-scale adjustment: compared to the food patch previously visited, spider monkeys traveled to larger patches just after fusions. This was not without an immediate travel cost: the interpatch distance and travel time after a fusion were longer than that before the fusion. This rapid adjustment shows the flexibility that fission–fusion dynamics can offer. Spider monkeys are in large subgroups only when food conditions are favorable, as evidenced by the fact that at all the other time-scale levels larger subgroups did not experience greater ranging costs than smaller subgroups. Our results indicate that on the whole spider monkeys successfully minimize ranging costs by fission and fusion of subgroups.     

The role of thermoregulation in lizard biology: Predatory efficiency in a temperate diurnal basker

Summary Decreasing levels of simulated solar radiation have the following effects in the diurnal basking lizard Lacerta vivipara: (i) increase in time spent basking with a consequent decrease in time available for foraging (ii) decrease in speed of movement whilst foraging (iii) decrease in total foraging distance, and hence contact with potential prey (iv) decrease in searching efficiency in an experimental arena. Complete absence of simulated solar radiation accentuates these effects, and reduces the proportion of faster-moving prey in the diet. Time taken to swallow prey (handling time) increases exponentially with decreasing body temperature. It is concluded from these results that maintaining relatively high activity temperatures (30–36°C) is adaptive for the species because the loss of potential foraging time caused by lengthy periods of basking is offset by the following advantages: increased contact with and capture of prey, increased efficiency of prey handling, and availability of a wider range of prey types.     

Group orientation in Neomysis mirabilis (Mysidacea: Crustacea)

The orientational behaviour of the neritic mysid Neomysis mirabilis in a swarm is considered under natural and experimental conditions. In nature, the swarms exhibit elements of inner integration, such as homogeneous age structure, collective behaviour, and relative constancy of inter-individual spacing. The mysids are guided by combined optomotor and vibrotactile cues, determining their disposition in aquaria of different shapes. Maximum accuracy of spatial arrangement is observed in round vessels. It is suggested that the stress encountered in an aquarium produces responses comparable to those elicited by the presence of a predator. The rigid spatial arrangement at the supra-individual level is interpreted as evolutionally-optimized prey strategy. These elements in the geometrical distribution of individuals in mysid swarms are inherent in other pelagic crustaceans also, and are not controlled by random taxes stimulated by hydrological factors.     

Thermal behaviour of a small spider (Araneae : Araneidae : Araneinae) on horizontal webs in semi-arid Western Australia

Summary The solar orientation response of a small spider (gen. nov.: Araneae: Araneidae: Araneinae) is examined at a semi-arid location in tropical Western Australia. The spiders rest beneath horizontal dome-shaped webs close to the ground surface on spinifex (Triodia sp.). The spiders are small (c. 5 mm long) and robust, being only twice as long as wide. Spider orientation, elevation, wind velocity, radiation and wet and dry bulb temperatures together with spider and egg-sac temperatures were measured throughout the day. The spiders orientated their long axis to the solar azimuth through much of the day and postured to the solar elevation; in so doing they tracked the apparent movement of the sun throughout the day with considerable precision, far exceeding that of other spiders studied. Stepwise regression indicated that the solar position was the most important environmental factor associated with the posturing and orientation. The variance in the orientation of the population was associated with a suite of environmental variables related to wind velocity and temperature. Spider (abdominal) temperature had a curvilinear relationship with ambient temperature, with a plateau at c. 35° C, whereas egg-sac temperature had a strictly linear relationship with ambient temperature. Stepwise regression indicates that both the spider and egg-sac temperatures are related mainly to net radiation and ambient temperature. The spiders can alter the incident radiation to which they are exposed by c. 2.2 times which represent the limits of the exposed silhouette area between full dorsal sunlight and the posterior (or anterior) aspect. Under hot conditions the spiders posture and orientate such that the long axis of the body is orientated abdomen towards the sun.     

Two clinal trends in male-male aggressiveness in a subsocial spider mite (Schizotetranychus miscanthi)

We studied the correlation between male-male aggressive behavior of the spider mite, Schizotetranychus miscanthi and winter coldness, a factor hypothesized to correlate with the degree of relatedness of the interacting males through the frequent occurrence of mother-son mating in the spring. Thirty-one samples from various populations throughout Japan were added to previous data. A cluster analysis revealed two clinal trends for male-male aggression in S. miscanthi in regions of light or no snowfall. The two clines may represent a dimorphism within the species, each having a similar clinal trend within Japan. Further support for the hypothesis that male overwintering survival affects male-male aggressiveness came from a comparison between regions of heavy snowfall and those of light or no snowfall. In regions with heavy snow insulation, overwintering survival may not be as high as in other areas, and there was no correlation between male aggressiveness and the minimum winter temperature. However, there was a significant correlation in regions of light snow. Received: 26 May 1998 / Received in revised form: 16 December 1998 / Accepted: 30 December 1998     

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     

Behavioral responses of a larval crustacean to hydrostatic pressure: Rhithropanopeus harrisii (Brachyura: Xanthidae)

Responses of the four zoeal stages of the crab Rhithropanopeus harrisii (Gould) to step and continuous changes in hydrostatic pressure were analyzed with a video system. Crabs were collected from the Neuse River estuary (North Carolina, USA) from June to August, 1987. The lower thresholds for step increases and decreases in pressure were 3 and 8 to 10 mbar, respectively. There was little change in sensitivity with zoeal development. Tests of larval responses in a light field that simulated the underwater angular light distribution indicated positive phototaxis does not occur upon pressure changes. In darkness, rates of pressure increase at and above 0.175 mbar s^-1 induced high barokinesis and negative geotaxis in all but Stage IV zoeae, which had a threshold of 1.19 mbar s^-1. Since larval sinking and descent swimming speeds exceed these threshold rates, larvae can move rapidly enough to produce suprathreshold changes in pressure which evoke behavioral responses. Slow rates of pressure decrease induced passive sinking while rapid rates caused an active ascent. This ascent response upon a pressure decrease is unreported among crustaceans, and is hypothesized to function for avoidance of feeding and respiratory currents of benthic invertebrates. The descent response occurs in all zoeal stages, except IV, at rates of pressure decrease (0.4 to 0.71 mbar s^-1) that are within the range of ascent swimming speeds. These results support Sulkin's negative feedback depth regulation model. The absolute distances moved before corrective vertical responses to threshold rates of pressure change are initated delimit the depth regulatory window. In darkness, the asymmetry of the window would lead to an ascent. It is hypothesized that light is an additional component in depth regulation, and that the limits and symmetry of the depth regulatory window may be controlled by the level of light adaptation.     

Eusociality and extraordinary sex ratios in the spider Anelosimus eximius (Araneae: Theridiidae)

Summary Colonies of Anelosimus eximius in Panama had an average sex ratio of 0.15±sd 0.09, i.e. about five females for each male. The sex ratio in egg sacs reared was even lower (0.08±0.01), as was that of immatures in newly founded colonies (0.12±0.05). The possible mechanisms responsible are discussed. Mature colonies had an average ratio of 17 females and 2 males for each egg sac present (range: 2–91 females, 0.2–8.2 males) and contained a large proportion of females which were not inseminated but which presumably help. Since both sexes are diploid, arrhenotoky can be ruled out and it is assumed that some females do not come to reproduction, the proportion depending on the availability of resources. This mechanism may enable entire colonies to survive lean times.     

Condition dependence of male display coloration in a jumping spider (Habronattus pyrrithrix)

In many animals, conspicuous coloration functions as a quality signal. Indicator models predict that such colors should be variable and condition dependent. In Habronattus pyrrithrix jumping spiders, females are inconspicuously colored, while males display brilliant red faces, green legs, and white pedipalps during courtship. We tested the predictions of the indicator model in a field study and found that male body condition was positively correlated with the size, hue, and red chroma of a male’s facial patch and negatively correlated with the brightness of his green legs. These traits were more condition dependent than non-display colors. We then tested a dietary mechanism for condition dependence using two experiments. To understand how juvenile diet affects the development of coloration, we reared juvenile spiders on high- and low-quality diets and measured coloration at maturity. To understand how adult diet affects the maintenance of coloration, we fed wild-caught adults with high- or low-quality diets and compared their coloration after 45 days. In the first experiment, males fed high-quality diet had redder faces, suggesting that condition dependence is mediated by juvenile diet. In the second experiment, red coloration did not differ between treatments, suggesting that adult diet is not important for maintaining the color after it is produced at maturity. Diet had no effect on green coloration in either experiment. Our results show different degrees of condition dependence for male display colors. Because red is dependent on juvenile diet, it may signal health or foraging ability. We discuss evidence that green coloration is age dependent and alternatives to indicator models for colorful displays in jumping spiders.     

Trade-off between pre- and postcopulatory sexual cannibalism in a wolf spider (Araneae,Lycosidae)

Sexual cannibalism can occur before, during or after mating. Relatively few experimental studies have examined why there is variation in the timing of sexual cannibalism. We examined the latency and number of attacks required for female spiders to capture male spiders pre- vs. postcopulation. We also examined the effects of female mating status and hunger level on the occurrence of pre- and postcopulatory cannibalism, which reflects the contribution of both relative capture success and female motivation to cannibalize males. Precopulatory cannibalism occurred after a shorter interval and required fewer chases and physical interactions for the female to successfully capture the male than was the case for postcopulatory cannibalism. Virgin females were more likely to engage in postcopulatory rather than precopulatory cannibalism and mated females vice versa. Those virgin females that did engage in precopulatory cannibalism had significantly lower body condition than virgin females engaging in postcopulatory cannibalism. While precopulatory cannibalism occurred more quickly and required fewer attacks by females, it comes at a potential cost of not mating with males. Hence, females are more likely to engage in precopulatory cannibalism if they have already mated or, if virgins, if they have low body condition. These results indicate that the decision of when to cannibalize males is dynamic and depends upon the relative value of a male as a mate versus a meal.     

Dispersal behaviour in a subsocial spider: group conflict and the effect of food availability

This study investigates proximate factors influencing dispersal behaviour in the subsocial spider Amaurobius ferox. Dispersal of spiderlings from the natal web occurred as a progressive sigmoidal function (mean duration of dispersal period=31 days), with considerable individual variability in developmental instar and body mass within the clutch at the time of dispersal. The spiderlings showed a significant decrease in group cohesion on the 6th day after their second moult, which corresponded to the beginning of the dispersal period. Mutual aggressiveness appeared when the spiderlings began to show predatory behaviour (4th day post-second moult) and increased over the course of the dispersal period. While lack of prey in the maternal nest accelerated dispersal behaviour, the addition of prey items lengthened the dispersal period in previously non-fed clutches. Individuals that dispersed were smaller than the remaining individuals when measured on the day 50% of the clutch had dispersed. Timing of the appearance of the developmental characters (second moult, predation activity, agonistic behaviour against siblings, reduction of group cohesion, dispersal) suggests that the dispersal trait might have evolved in consequence of these different functional behaviours. Received: 10 August 1999 / Received in revised form: 15 March 2000 / Accepted: 1 April 2000     

Arsenic (As), antimony (Sb), and lead (Pb) availability from Au-mine Technosols: a case study of transfer to natural vegetation cover in temperate climates

Soils from old Au-mine tailings (La Petite Faye, France) were investigated in relation to the natural vegetation cover to evaluate the risk of metals and metalloids (Pb, As, Sb) mobilizing and their potential transfer to native plants (Graminea, Betula pendula, Pteridium aquilinum, Equisetum telmateia). The soils are classified as Technosols with high contamination levels of As, Pb, and Sb. The single selective extractions tested to evaluate available fraction (CaCl₂, acetic acid, A-Rhizo, and DTPA) showed low labile fractions (<5 % of bulk soil contents), but still significant levels were observed (up to 342.6 and 391.9 mg/kg for As and Pb, respectively) due to the high contamination levels of soils. Even at high soil contaminations (considered as phytotoxic levels for plants), translocation factors for native plants studied are very low resulting in low concentrations of As, Sb, and Pb in their aerial part tissues. This study demonstrates the important role of (1) native plant cover in terms of “stabilization” of these contaminants, and (2) the poor effectiveness of extraction procedures used for this type of soil assemblages, i.e., rich in specific mineral phases.