Cross-modality effects during male-male interactions of jumping spiders

An important prediction from game theory is that the value of a resource influences the level to which male-male conflict escalates. Earlier experimental studies have shown that the seven salticid species we study here (Bavia aericeps, Euryattus sp., Hypoblemum albovittatum, Jacksonoides queenslandicus, Marpissa marina, Portia africana and Simaetha paetula) determine by sight whether a female is a conspecific or a heterospecific and then escalate the intensity with which they interact (i.e., they adopt behaviour that is likely to put them at greater risk of injury after detecting the presence of a conspecific female). Here the earlier studies are extended by using the odour of conspecific females (experimental tests) and heterospecific females (control tests), and by presenting each male with his mirror image as well as having two males interact with each other. Findings from this study suggest that, for J. queenslandicus and P. africana, the odour of conspecific females, more than the odour of heterospecific females, primes the male to escalate conflict with a potential rival. However, this was not found for the other five species tested.     

Influence of seeing a female on the male–male interactions of a jumping spider, Hypoblemum albovittatum

An important prediction from game theory is that resource value influences the level to which competitors will escalate conflict. An earlier study considered whether this prediction applies to the male–male interactions of Hypoblemum albovittatum, a jumping spider (Salticidae) from New Zealand. The males of this species escalated conflicts in the presence of a moving mount made from a conspecific female. However, because the control was only a similar-sized motionless cork, an alternative hypothesis (that the cue for escalation is seeing movement of any female-size object, rather than seeing specifically a female) was not ruled out. Here we show that a moving cork, without a mount present, is indeed sufficient to cause males to escalate, but a moving mount (made from a conspecific female) causes males to escalate further. The level of escalation in the presence of a moving mount made from prey (housefly) or from a rival (conspecific male) did not differ significantly from the level of escalation when only a moving cork was present. These findings suggest that, although seeing a moving object similar in size to a conspecific female is sufficient for priming males to escalate, males can also discern by sight that specifically a female is present and, when they have this precise information, they make strategic decisions to escalate conflict further.     

Effects of prey–spider odour on intraspecific interactions of araneophagic jumping spiders

An important prediction from game theory is that the value of a resource will influence the level to which an animal is willing to escalate during conflict with conspecific rivals. Here we use this prediction as the rationale for experiments aimed at determining whether escalation decisions made by predators are influenced by the presence of preferred prey. The predators we use are adult individuals of Portia fimbriata, a jumping-spider (Salticidae) species from Queensland (Australia). P. fimbriata is known to prefer other salticid species as prey and can detect the odour of Jacksonoides queenslandicus, this being an especially common salticid species in the Queensland habitat. Here we show that, for P. fimbriata females, the odour of J. queenslandicus primes escalation of vision-based conflict. However, for P. fimbriata males, no comparable effect was found. These findings suggest intrasexual competition for prey is more important to females than to males.     

How cross-modality effects during intraspecific interactions of jumping spiders differ depending on whether a female-choice or mutual-choice mating system is adopted

An important prediction from game theory is that the value of a resource influences the level to which conflict escalates. Here we use jumping spiders (Salticidae) to consider this prediction in the context of species adopting different mating systems (‘female mate-choice’ and ‘mutual mate-choice’). Our experiments are designed for determining whether the odour of conspecific females, more than the odour of heterospecific females, primes males to escalate conflict with a potential same-sex rival and also whether the odour of conspecific males, more than the odour of heterospecific males, primes females to escalate conflict with a potential same-sex rival. Four species were studied: Evarcha culicivora, a species in which mutual mate-choice is pronounced, and Portia fimbriata, Portia africana, and Jacksonoides queenslandicus, more conventional salticids in which female mate-choice and male–male competition appear to be dominant. Our hypothesis is that, for all four species, there is strong competition between males for access to females and that, for E. culicivora, but not for the other three species, there is also strong competition between females for access to males. Our findings are consistent with this hypothesis, as we show that, although the odour of conspecific females primes escalation of vision-based male–male conflict for all four species, E. culicivora is the only species for which there is evidence of odour from conspecific males priming the escalation of female–female conflict.     

Interspecific differences in the movements of female ticks on reptiles.

Movements of females of two reptile ticks, Amblyomma limbatum and Aponomma hydrosauri were studied to determine whether the behaviour of the species on hosts was similar or different. Experiments were conducted to determine if the movement of females on hosts prior to mating was influenced by the delayed attachment of conspecific males. A field experiment was also undertaken to determine whether data obtained during laboratory experiments reflected the behaviour of females on hosts in the field. Both in laboratory and field experiments, a significantly greater proportion of Ap. hydrosauri females changed their site of attachment prior to mating. Individual Ap. hydrosauri females made significantly more positional changes than Amb. limbatum females. These interspecific differences occurred irrespective of the absence of conspecific males or presence of non-conspecific adults on hosts. These data may have important ecological implications in relation to the success of colonizing females at their parapatric boundary.     

Cues by which suspended-leaf nests of Euryattus (Araneae: Salticidae) females are recognized by conspecific males and by an aggressive-mimic salticid, Portia fimbriata

Females of Euryattus sp. indet, a salticid from Queensland, suspend rolled-up leaves for nests. Euryattus males respond to conspecific females in nests with vibratory courtship. Portia fimbriata , a sympatric salticid that preys on Euryattus , responds to Euryattus females' nests by mimicking the courtship of Euryattus males. In the laboratory, cues by which potential mates (conspecific males) and potential predators ( P. fimbriata females) recognize the nests of Euryattus females are investigated experimentally. Both the position of the leaf in relation to the substratum and the number of threads between the leaf and the substratum appear to be important cues by which both the Euryattus males and P. fimbriata females find nests. In addition, chemical cues from Euryattus females influence conspecific males, but there is no evidence that chemical cues are important to the predator.     

Reproductive interference between three parapatric species of reptile tick

Summary Reproductive interference between three species of reptile tick, Aponomma hydrosauri, Amblyomma albolimbatum and Amb. limbatum was investigated. Adults of two species attach together on the same lizard hosts in narrow overlap zones at parapatric boundaries between species, providing opportunities for interference. The possibility of reproductive interference was suggested because of similarities in the sexual communication systems.Three forms of interference were found in laboratory experiments. Firstly male movement to search for sexually receptive females was inhibited when females of two species were present on the same host, probably because of signal interference. Secondly, a nonspecific, short range attractant pheromone led males into non-conspecific courtship attempts which reduced the time for conspecific courtships. Thirdly Amb. albolimbatum males physically blocked the genitalia of female Ap. hydrosauri after unsuccessful non-conspecific courtship attempts. Field data showed this behaviour was common in both Amblyomma species after conspecific matings. Reproductive interference would reduce the fitness of each species in sympatry, and may contribute to the maintenance of the narrow parapatric boundaries.     

Widespread reliance on olfactory sex and species identification by lyssomanine and spartaeine jumping spiders

Jumping spiders (Salticidae) are renowned for their exceptional vision, but this does not preclude use of other senses. Here we provide evidence that olfactory pheromones are widespread in the Spartaeinae and Lyssomaninae, two subfamilies regarded as basal clades within the Salticidae. Pheromone use by salticids was tested in a series of experiments: males were tested with the odour of conspecific females, heterospecific females, and conspecific males, and females were tested with the odour of conspecific males. With seven of the 29 species tested, we also tested males using the draglines of conspecific females (spider absent) as the odour source. Males of all species tested were attracted to the odour of conspecific females and to the odour of the draglines of conspecific females. There was no evidence of males responding to the odour of heterospecific females or conspecific males, or of females responding to the odour of conspecific males. These findings suggest that it is primarily males that respond to olfactory sex pheromones, consistent with the apparent trend within spiders of males more actively searching for females and females placing greater emphasis on mate‐choice decisions. Compared with most salticid groups, lyssomanines and spartaeines are unusually sedentary and this lifestyle may favour olfactory mate searching. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 664–677.     

Preference for conspecifics evolves earlier in males than females in a sexually dimorphic radiation of fishes

Speciation by sexual selection is generally modeled as the coevolution of female preferences and elaborate male ornaments leading to behavioral (sexual) reproductive isolation. One prediction of these models is that female preference for conspecific males should evolve earlier than male preference for conspecific females in sexually dimorphic species with male ornaments. We tested that prediction in darters, a diverse group of freshwater fishes with sexually dimorphic ornamentation. Focusing on the earliest stages of divergence, we tested preference for conspecific mates in males and females of seven closely related species pairs. Contrary to expectation, male preference for conspecific females was significantly greater than female preference for conspecific males. Males in four of the 14 species significantly preferred conspecific females; whereas, females in no species significantly preferred conspecific males. Relationships between the strength of preference for conspecifics and genetic distance revealed no difference in slope between males and females, but a significant difference in intercept, also suggesting that male preference evolves earlier than females’. Our results are consistent with other recent studies in darters and suggest that the coevolution of female preferences and male ornaments may not best explain the earliest stages of behavioral isolation in this lineage.     

The biology of ant-like jumping spiders: intraspecific interactions of Myrmarachne lupata (Araneae, Salticidae)

Myrmarachne lupata is an ant-like salticid in which males have very large chelicerae. The display repertoire of this species is unusually large and complex for a salticid spider. Each individual male uses one of three different mating tactics depending on the female's maturity and location. With adult females outside nests type 1 courtship occurs which seems to be a form of visual communication and includes specialized movements and postures of the legs, palps and body. With adult females inside nests, males use type 2 courtship, which seems to be a form of non-visual communication and consists primarily of probing with the legs on the silk; males mate with receptive females inside the nests. With subadult females, males first use type 2 courtship then spin an adjacent silken chamber and cohabit. After she moults and matures, mating occurs inside the nest. Vacant nests of conspecific females, but not those of another sympatric salticid species, elicit courtship behaviour from males. During male-male interactions, embracing occurs with the large chelicerae spread apart. Females and subadults also display, and different displays occur in interactions depending on the sex/age classes of the spiders involved. Despite the unusual morphology of these spiders, their individual displays are similar to those of more typical salticids. During copulation males stand beside the female instead of over or on her as occurs with typical salticids.     

Sex ratio response of the parasitoid wasp Muscidifurax raptor to other females

This study examines the sex ratio response of the parasitoid wasp Muscidifurax raptor to conspecific and confamilial females in relation to two groups of functional sex ratio models, local mate competition and host quality models. In some but not all experiments, M. raptor females produced a greater proportion of sons in the presence of a conspecific female than when alone, and this sex ratio effect carried over for a day after the females were isolated from each other M. raptor females also produced a greater proportion of sons in the presence of a female of the confamilial parasitoid Spalangia cameroni than when alone (although only on the second day of exposure to S. cameroni, not on the first). M. raptor's sex ratio increase in the presence of conspecifics is consistent with local mate competition models but not with host quality models because the presence of a conspecific female did not cause there to be more, and thus potentially smaller, offspring developing per host. In contrast, the presence of a S. cameroni female did cause there to be more offspring developing per host than when a M. raptor female was alone; thus M. raptor's sex ratio increase in the presence of S. cameroni may be explained by host quality models. An alternative explanation for the sex ratio increase in response to confamilials is that only a sex ratio response to conspecifics may be adaptive, due to local mate competition; but M. raptor females may be unable to distinguish between conspecific and S. cameroni females.     

Mating behaviour and reproductive isolation of three species of reptile tick

The mating behaviour of three species of reptile tick, Aponomma hydrosauri, Amblyomma albolimbatum and Amb. limbatum is similar and involves a female sex pheromone which activates males to detach from their hosts and search. After contacting females, a stereotyped six-phase courtship sequence occurs. There are qualitative differences between the species in courtship behaviour at phase 3 (reversal of position by dorsally mounted males) and phase 6 (the copulation position). On-host observations of non-conspecific mating show that females of the three species have species-specific activation pheromones, which is contrary to reports in other species of tick. Such specificity should result in reproductive isolation of the three species; however, under certain circumstances it may not prevent non-conspecific contacts between the sexes. Off-host observations of courtship behaviour show that once males of the three species contact non-conspecific females, they attempt courtship and are persistent with their courtship. Rarely did non-conspecific courtships proceed beyond phase 4 of the courtship sequence, as non-conspecific females did not lift their bodies to allow males venter contact. Differences between the species in leg orientation in the copulation position, together with body size differences, are responsible for a complete barrier to successful non-conspecific copulation. The observations illustrate the role that behavioural mechanisms play in reproductively isolating these three species of tick.     

Female Japanese quail aggregate to avoid sexual harassment by conspecific males: a possible cause of conspecific cueing

In a series of three experiments, we examined the hypothesis that female Japanese quail, Coturnix japonica, are motivated to form aggregations to reduce the probability of harassment by conspecific males. When in the presence of a conspecific male, female quail both sought others of their sex and remained near them. However, if no males were present, females were indifferent to or tended to avoid one another. We interpret these data as consistent with the hypothesis that one function of aggregations of female quail is to avoid harassment by conspecific males. Skew in male mating success would be a probable consequence of such aggregations of females. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Love is in the air: olfaction-based mate-odour identification by jumping spiders from the genus Cyrba

Jumping spiders (Salticidae) are known for having good eyesight, but the extent to which they rely on olfaction is poorly understood. Here we demonstrate for the first time that olfactory pheromones are used by two species from the salticid genus Cyrba (C. algerina and C. ocellata). Using a Y-shape olfactometer, we investigated the ability of adult males and females of both species to discriminate between mate and non-mate odour. A hidden spider or a spider’s draglines (no spider present) were used as odour sources. There was no evident response by females of either Cyrba species to any tested odour. Males of both species chose odour from conspecific females, or their draglines, significantly more often than the no-odour control, but there was no evident response by males to any of the other odours (conspecific male and heterospecific female). Our findings demonstrate that C. algerina and C. ocellata males can make sex- and species-specific discriminations even when restricted to using olfaction alone. Also, by showing that draglines can be a source of olfactory pheromones, our findings illustrate the difficulty of ruling out olfaction when attempting to test for chemotactile cues.     

Sex pheromones in maleMelittobia parasitic wasps: Female response to conspecific and congeneric males of 3 species

Presented a choice between conspecific males and 2 congeneric males, virgin females ofMelittobia australica andM. digitata chose conspecific males disproportionately more often, whereasM. femorata females distributed themselves evenly among the choices. Empty tubes, provided as the fourth choice in the test apparatus, were entered much less often than tubes containing live males. Females of all species chose “wrong” males about equally frequently. These observations suggest that even non-conspecific males possess some degree of attractiveness to virgin females. Chemicals in the sex pheromone of the males are presumed to be the source of the males' attractancy. The incomplete species specificity is interpreted in light of the life history of this genus, and it is suggested that specific recognition cues operate primarily after the sexes come together. Supported by a grant from Kagoshima Prefecture in Japan under the exchange program of faculty members between Kagoshima University and the University of Georgia.     

Prey Habitat Location by the Cassava Mealybug Predator Exochomus flaviventris: Olfactory Responses to Odor of Plant, Mealybug, Plant–Mealybug Complex, and Plant–Mealybug–Natural Enemy Complex

Exochomus flaviventris Mader is considered to be the most active predator of the cassava mealybug Phenacoccus manihoti Matile–Ferrero in Central Africa. The response of experienced gravid female coccinellids to the odor of cassava plant (var. Zanaga), unparasitized mealybugs, plant–mealybug complex with or without feeding prey (parasitized or not), and plant–mealybug complex with or without conspecific coccinellids was investigated in a Y-tube olfactometer. The odor of uninfested cassava plants was not more attractive than clean air. Dual-choice tests revealed that mealybug-infested plants were preferred to mealybugs alone and mealybug-damaged plants and were the major sources of volatiles that attract females coccinellids to the microhabitat of its prey. The emission of volatile chemicals did not appear to be limited to the infested parts of the plant but did occur systemically throughout the plant. The presence of conspecific coccinellid larvae or adult males did not modify the attractiveness of the mealybug-infested plants. However, when an infested plant with conspecific predator females (alone or with conspecific males) was compared to an infested plant or infested plant with conspecific males, E. flaviventris females showed a preference for the last two sources of odor. The uninfested plant with conspecific males was also preferred to the uninfested plant with conspecific females. In addition, the odor of conspecific males was preferred over that of conspecific females. Female predators preferred the plant infested with unparasitized mealybugs over the plant infested with mealybugs previously parasitized. These results showed that E. flaviventris females use herbivore-induced plant volatiles during foraging and can detect via olfaction the presence of conspecific gravid females and parasitized prey, thus assessing patch suitability from a distance.     

Release of chemicals by prostaglandin-treated female fathead minnows, Pimephales promelas, that stimulate male courtship

Intramuscular injection of PGF2 alpha tromethamine salt (Sigma) did not induce sexual behavior in female fathead minnows, Pimephales promelas, when in the presence of male conspecific. However, coresident males exhibited an increased frequency of courtship behavior, consisting of approach and leading, to females following PGF2 alpha treatment. When presented with both saline-injected and PGF2 alpha-injected females, males showed increased courtship only to PGF2 alpha-treated females. Isolated males also showed increased leading behavior following the introduction of water exposed to PGF2 alpha-treated females, but not with water exposed to PGF2 alpha-treated males, untreated females or males, or PGF2 alpha alone. Hence, PGF2 alpha appears to induce the release of a female-specific chemical in P. promelas that triggers courting behavior in conspecific males.     

The Functioning of Species-Specific Olfactory Pheromones in the Biology of a Mosquito-Eating Jumping Spider from East Africa

Having unique, complex eyes and vision based on exceptional spatial acuity, salticid spiders are known for their vision-based predatory and mating strategies. Yet Evarcha culicivora, the salticid we consider here, is known for predatory and mating strategies based strongly on the interplay of vision and olfaction. This unusual East African species feeds indirectly on vertebrate blood by choosing blood-carrying female mosquitoes as preferred prey, which it can identify by sight and by smell. Moreover, E. culicivora’s mating strategy is unusual because, unlike the prevailing general pattern among spiders, where males are more active in courtship and females are more active in mate choice, both roles are characteristic of both sexes of E. culicivora. There is also an unusual relationship between diet and mating in this species, with blood meals making both sexes more attractive as potential mates. However, findings from the present study demonstrate that, regardless of source-spider diet and even when restricted to using chemoreception in the absence of seeing another spider, both sexes can discriminate between opposite-sex conspecifics and opposite-sex heterospecifics. Yet there is no evidence that diet (blood versus no blood) influences the attractiveness of opposite-sex heterospecific individuals to E. culicivora. Evidence that the odor of opposite-sex conspecifics is salient to E. culicivora comes from three different experimental designs (retention testing, choice testing and courtship-initiation testing). The effective odor source can be either the presence of the spider or the presence of the spider’s draglines alone.     

Femme Fatale — the Case of the Threespine Stickleback

In addition to courting females, male threespine sticklebacks must guard their nests from attacks on the eggs by female cannibals. We hypothesized that the presence of females in high densities in tide pools adversely affects a male's health and reproductive success. In the laboratory males housed with conspecific females during the breeding season were in poorer physical condition and died sooner than solitary males and males housed with conspecific males or with heterospecific females.     

Intraspecific acoustic communication and mechanical sensitivity of the tympanal ear of the gypsy moth Lymantria dispar

Tympanal ears of female gypsy moths Lymantria dispar dispar (L.) (Lepidoptera: Erebidae: Lymantriinae) are reportedly more sensitive than ears of conspecific males to sounds below 20 kHz. The hypothesis is tested that this differential sensitivity is a result of sex‐specific functional roles of sound during sexual communication, with males sending and females receiving acoustic signals. Analyses of sounds produced by flying males reveal a 33‐Hz wing beat frequency and 14‐kHz associated clicks, which remain unchanged in the presence of female sex pheromone. Females exposed to playback sounds of flying conspecific males respond with wing raising, fluttering and walking, generating distinctive visual signals that may be utilized by mate‐seeking males at close range. By contrast, females exposed to playback sounds of flying heterospecific males (Lymantria fumida Butler) do not exhibit the above behavioural responses. Laser Doppler vibrometry reveals that female tympana are particularly sensitive to frequencies in the range produced by flying conspecific males, including the 33‐Hz wing beat frequency, as well as the 7‐kHz fundamental frequency and 14‐kHz dominant frequency of associated clicks. These results support the hypothesis that the female L. dispar ear is tuned to sounds of flying conspecific males. Based on previous findings and the data of the present study, sexual communication in L. dispar appears to proceed as: (i) females emitting sex pheromone that attracts males; (ii) males flying toward calling females; and (iii) sound signals from flying males at close range inducing movement in females, which, in turn, provides visual signals that could orient males toward females.