The Elaborate Postural Display of Courting <Emphasis Type="Italic">Drosophila persimilis</Emphasis> Flies Produces Substrate-Borne Vibratory Signals

Sexual selection has led to the evolution of extraordinary and elaborate male courtship behaviors across taxa, including mammals and birds, as well as some species of flies. Drosophila persimilis flies perform complex courtship behaviors found in most Drosophila species, which consist of visual, air-borne, gustatory and olfactory cues. In addition, Drosophila persimilis courting males also perform an elaborate postural display that is not found in most other Drosophila species. This postural display includes an upwards contortion of their abdomen, specialized movements of the head and forelegs, raising both wings into a “wing-posture” and, most remarkably, the males proffer the female a regurgitated droplet. Here, we use high-resolution imaging, laser vibrometry and air-borne acoustic recordings to analyse this postural display to ask which signals may promote copulation. Surprisingly, we find that no air-borne signals are generated during the display. We show, however, that the abdomen tremulates to generate substrate-borne vibratory signals, which correlate with the female’s immobility before she feeds onto the droplet and accepts copulation.     

Infestation with the parasitic nematode <Emphasis Type="Italic">Philometra ovata</Emphasis> does not impair behavioral sexual competitiveness or odor attractiveness of the male European minnow (<Emphasis Type="Italic">Phoxinus phoxinus</Emphasis>)

Our understanding on the role of chemical signals in parasite-mediated sexual selection is still limited, and only some existing studies have focused on fish. Furthermore, published studies on the effect of parasite infection on behavioral sexual competition of the male hosts have yielded contradictory results. Here, we examined whether the infection of the body cavity-dwelling parasitic nematode Philometra ovata influences odor-based female choice and behavioral sexual competition (dominance and courtship behavior) between males in the European minnow (Phoxinus phoxinus), the cyprinid fish host. In contrast to our predictions, we found that naïve females showed no preference between the odors of infected and non-infected males, thus indicating that P. ovata infection may not affect odor-based female choice. Moreover, P. ovata did not impair sexual competitiveness of their hosts either. Our results indicate that despite its relatively large size, P. ovata may not alter sexual cues and the success of the male hosts in sexual selection.     

The Role of Males in Host-Fruit Selection by Females of a Walnut Infesting Tephritid (Diptera) <Emphasis Type="Italic">Rhagoletis zoqui</Emphasis>

Rhagoletis zoqui Bush flies have a mating system in which males guard and defend walnut fruit-hosts from other males and mate, apparently without courtship, with females as they arrive to oviposit. Hypothetically, female selection of a particular fruit may be due to the quality of fruit for larval development (previously determined by guarding males), the quality of the male upon the fruit as a mate, or both, but this is not clear. We performed an experiment to determine if R. zoqui females or males select the fruit to oviposit or guard based on its quality for larval development (i.e., size, sugar-content [brix] or hardness), or following male-mediated cues such as chemical residues, related to prior fruit occupation by the male during guarding. Fruit choice by R. zoqui females and males were examined under semi-natural conditions on caged branches of English walnut trees, Juglans regia L., growing in the highlands of south-central Mexico. A single male or female was allowed to select a fruit and was then removed. An individual of the opposite sex was then introduced to the same branch and presented with the opportunity to choose among the same array of hosts. The pattern of fruit choice was consistent, with female choice on the basis of male presence, suggesting that males left a chemical cue that persisted in their absence. No fruit quality differences were detected between selected and non-selected fruit. Fruit features selected by females and males matched almost exactly, although males rarely select the same fruit as females. We conclude that females preferentially selected fruit previously occupied by males although additional studies are required to determine the cues used by R. zoqui for oviposition resource selection by both females and resource guarding males.     

Female preference for both behavior and morphology traits of the male Japanese newt <Emphasis Type="Italic">Cynops pyrrhogaster</Emphasis>

Males of the Japanese newt (Cynops pyrrhogaster) display complex courtship behaviors and are sexually dimorphic in terms of tail morphology. Pair encounter experiments were conducted to investigate which males are preferred by female newts. Male courtship behavior consisted of four stages, namely, Approach, Fan, Creep and Spermatophore deposition. The Fan behavior was classified into four sub-patterns. Males which showed a specific sub-pattern were accepted at a significantly higher probability by females than males which showed the other sub-patterns. The accepted males had a smaller snout–vent length, higher tail, and larger body mass than the rejected males, and their body weight was relatively heavier. Our results suggest that females of C. pyrrhogaster select their mates based on both behavioral patterns and morphological characters.     

Social Relationships between Males and Females in Rock Lizard (<Emphasis Type="Italic">Darevskia brauneri</Emphasis>, Lacertidae). 2. Searching for an Area of Stable Residence,Criteria for Choice of Social Partner,and Factors of Friendly Integration of Dyads

Long-termed intersexual friendly relationships between males and females of the Brauner’s lizard are characterized by high levels of spatial association and high frequency of affiliative behavior between partners (see report 1). In present article we consider changes in the frequency of affiliative behavior in males (Am) and females (Af) after the first appearance of female on the male’s territory and conditions for the transition of lizard to stable residency (SR). Generally, Am becomes stable during the first days of initial period and does not change later. This fact suggests that in most instances male’s choice criteria are some peculiarities of the female’s exterior. In the initial period, Af is always low, however, gradually rises to the level of Am if Am is high. The main condition of female’s SR is high Am of the owner of the territory where female establishes her comfort areas. Hence, social partner choice in female is based on male’s behavior, and high Am triggers the formation of friendly interrelations. Significant correlation between social and sexual interrelations is absent. As can be judged by female’s reactions to male’s sexual behavior, his sexual activity is rather a factor of social disintegration. Structure of costs for social partner search is different in males and females, and these differences well harmonize with differences in gender strategies of partner choice. The social strategies are well interpreted in terms of the optimal foraging theory.     

Stridulation and Courtship Behaviour of Four Endemic Pill-Millipedes, <Emphasis Type="Italic">Arthrosphaera</Emphasis> spp. (Sphaerotheriida: Arthrosphaeridae) of the Western Ghats of India

Mating behaviour of four species of pill-millipedes under genus Arthrosphaera Pocock (Arthrosphaera dalyi Pocock and Arthrosphaera disticta Pocock, Arthrosphaera fumosa Pocock and Arthrosphaera magna Attems) endemic to the Western Ghats of Southern India was analyzed in mesocosms. Stridulation is a classical communication signal in males as well as females for mate selection. Conglobation (or volvation) is a mechanism of defence to protect from disturbance or avoid predation. If male touches female or vice versa they conglobate. To avoid disparity among individuals of the same species, volvating pill-millipedes evolved stridulation behaviour for communication. The male broadcasts appropriate signal to female through stridulation to advertise its interest in mating. The females test the male’s fitness by conglobation and suitable male uncoils the partner through stridulation signals. Male with its pygidium successfully uncoils the female and attains suitable orientation for courtship. Male pairs with female ventro–ventro contact in opposite direction to deposit sperm into the vulva of female. The duration of mating varies from species to species and usually a lapse from 3 to 30 min. Vibration generated by stridulation is species-specific and its perception mechanism in pill-millipedes is yet to be clearly understood. Present study emphasized the structure of stridulatory organs, mechanism of stridulation and pattern of mating behaviour in four species of pill-millipedes.     

Abdomen Posture and Flash Gesture Direction in a Female Synchronic Firefly <Emphasis Type="Italic">Photinus carolinus</Emphasis> (Green) (Coleoptera: Lampyridae)

North American Photinus fireflies use bioluminescent flashes to communicate an individual’s species and sex, and to attract potential mates. A female firefly responds to a male firefly’s courtship flash with her own species-specific flash. We used a photic stimulator to produce male-like species-specific P. carolinus LED courtship flashes. These evoked species-specific response flashes from a female. The female’s flashes were preceded by a flash gesture comprising a sequence of abdominal postural adjustments (pitch, roll, and yaw). These gestures changed her lantern’s orientation which, at rest, was downward towards the substrate. Our results demonstrate that these gestures mediate a lateralization of the female’s response flashes towards the direction of the stimulating LED. That is, she directs her response to the left of midline when stimuli are presented from her left, and similarly, she directs her response to the right of midline when stimuli are presented from her right. The directional aspect of the flash gesture adds a new perspective to the complexity of the behaviors associated with flash communication in fireflies. Lateralization of the flash gesture suggests that the female’s visual system processes information about the location of male’s flashes as well as their temporal pattern.     

Courtship in Two Morphotypes of the <Emphasis Type="Italic">Anastrepha fraterculus</Emphasis> (Diptera: Tephritidae) Cryptic Species Complex and their Implications for Understanding Mate Recognition

Ritualized courtship behaviors are used to recognize potential mates and behavioral patterns are inevitably different among populations that demonstrate reproductive incompatibility. We characterized and compared the courtship behaviors of two morphotypes of the cryptic species complex Anastrepha fraterculus: Brazil-1 morphotype and Brazil-3 morphotype. Courtship behaviors were filmed to analyze the behavioral sequences of these two morphotypes during homotypic crossings. The behavioral units Alignment (AL) and Abdominal movements (AB and AB-call) were newly recognized in the courtship ethogram of Anastrepha fraterculus males. The two morphotypes show distinct behavioral sequences leading up to copulation. Some behaviors were repeated frequently during the courtship process, while others were more restricted to the final moments of courtship. The three behavioral units that contributed most to copulation success were Contact, Alignment, and Arrowhead 1 in the Brazil-1 morphotype and Alignment, Arrowhead 1, and Fanning in the Brazil-3 morphotype. Some behavioral routines differed across the two morphotypes. Significant differences were also noted between the frequencies of the behavioral units displayed during courtship in the two morphotypes. The relationships between the pre-zygotic incompatibilities of the Brazil-1 and Brazil-3 morphotypes and the differences between the courtship behaviors of their males are discussed. Our results indicate that behavioral isolation is involved in the process of pre-zygotic reproductive isolation of Brazil-1 and Brazil-3 morphotypes.     

Mating and aggressive behaviour of <Emphasis Type="Italic">Brachyrhaphis olomina</Emphasis> (Cyprinodontiformes: Poeciliidae)

Despite the increasing interest in the use of intromittent male genitalia and coercive mating behaviour in poeciliids, detailed studies of the mating behaviour of most species in this family are lacking. We describe here the mating and aggressive behaviours of Brachyrhaphis olomina, and correlate them with the condition of the female’s ovum and embryos (immature, mature and pregnant). B. olomina performed a wide range of aggressive (sidle spread, tail beating, coordinate) and mating behaviours (approximation, touch, lateral display, touch-lateral display). Some behaviours (e.g. tail beating) are shared with other poeciliids, but two sexual behaviours (touch and lateral display) and one aggressive (coordinate) behaviour may be unique to B. olomina and were not reported in a previous study. Differences in male behaviour when paired with a female with mature ovum (more mating displays, no agonistic movements) suggest that males detect the female’s reproductive condition from some distance. The distinctive nature of mating behaviour in B. olomina highlights the importance of studying different species to have a better understanding of the evolution of mating and aggressive behaviours in poeciliids. Digital video images related to the article are available at http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo01a, http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo02a and http://www.momo-p.com/showdetail-e.php?movieid=momo170720bo03a.     

Exploring the influence of individual courtship behaviors on male mating success in a blow fly

Male courtship behavior and displays influence female mating decisions, and therefore affect mating success in a diverse range of organisms. While there is substantial evidence confirming that females prefer males who invest more in courtship, less attention has been paid to the relative importance of individual behaviors, and the discrete sequences of courtship that result in mating success. The small hairy maggot blow fly, Chrysomya varipes, performs stereotyped courtship behaviors, involving orienting, tapping, waving, arching, wing vibration and mounting. This study aimed to quantify male investment in specific courtship behaviors, and compare courtship investment and behavioral transitions between males who gained mating success (successful males) and those that did not (unsuccessful males). Our results show that mating success was influenced by the behaviors orienting, tapping, arching and mounting. Behavioral transitions revealed a distinct pattern of behaviors leading to a mating attempt, and some differences were observed between successful and unsuccessful males. Overall, our findings suggest that female mating decisions were based on differences in specific male courtship behaviors. This detailed observational study has quantified multiple courtship behaviors for the first time in C. varipes, and highlights the importance of considering multiple behavioral traits when exploring the influence of male courtship on mating success.     

Female preference for large waving claws in the dotillid crab <Emphasis Type="Italic">Ilyoplax pusilla</Emphasis>

Male Ilyoplax pusilla perform a waving display, a simple up-down movement of the claws during the reproductive season. Large males dedicated most of their surface activity to waving displays and gained higher mating success. On the other hand, small males infrequently performed waving displays and devoted their time exclusively to foraging. To examine female preference for the size of waving males, two female-release experiments were conducted. In the experiments, we recorded female choice between small- and large-waving claw models over short (10 cm) and long (25 cm) distances. In both the short- and long-distance choice experiments, significantly more females chose the large-claw model over the small-claw model. The following characteristics may produce obvious age-dependent sexual advertisement in I. pusilla, which grows throughout a life: (1) the strong female preference for large claws; (2) the short, 2-year lifespan that includes only two reproductive seasons; and (3) the lack of alternative mating strategies (e.g., surface mating).     

Sexual dimorphism of sonic apparatus and extreme intersexual variation of sounds in <Emphasis Type="Italic">Ophidion rochei</Emphasis> (Ophidiidae): first evidence of a tight relationship between morphology and sound characteristics in Ophidiidae

Background

Many Ophidiidae are active in dark environments and display complex sonic apparatus morphologies. However, sound recordings are scarce and little is known about acoustic communication in this family. This paper focuses on Ophidion rochei which is known to display an important sexual dimorphism in swimbladder and anterior skeleton. The aims of this study were to compare the sound producing morphology, and the resulting sounds in juveniles, females and males of O. rochei.

Results

Males, females, and juveniles possessed different morphotypes. Females and juveniles contrasted with males because they possessed dramatic differences in morphology of their sonic muscles, swimbladder, supraoccipital crest, and first vertebrae and associated ribs. Further, they lacked the ‘rocker bone’ typically found in males. Sounds from each morphotype were highly divergent. Males generally produced non harmonic, multiple-pulsed sounds that lasted for several seconds (3.5?±?1.3 s) with a pulse period of ca. 100 ms. Juvenile and female sounds were recorded for the first time in ophidiids. Female sounds were harmonic, had shorter pulse period (±3.7 ms), and never exceeded a few dozen milliseconds (18?±?11 ms). Moreover, unlike male sounds, female sounds did not have alternating long and short pulse periods. Juvenile sounds were weaker but appear to be similar to female sounds.

Conclusions

Although it is not possible to distinguish externally male from female in O. rochei, they show a sonic apparatus and sounds that are dramatically different. This difference is likely due to their nocturnal habits that may have favored the evolution of internal secondary sexual characters that help to distinguish males from females and that could facilitate mate choice by females. Moreover, the comparison of different morphotypes in this study shows that these morphological differences result from a peramorphosis that takes place during the development of the gonads.

     

Courtship behavior,communicative sound production and resistance to stress in Drosophila mutants with defective <Emphasis Type="Italic">agnostic</Emphasis> gene coding for LIMK1

To elucidate the role of one of the main elements of signal cascade of actin remodeling—LIM-kinase 1 (LIMK1)—in the control of animal behavior we have studied the characteristics of courtship behavior, parameters of acoustic communicative signals and their resistance to heat shock (HS, 37°C, 30 min) in Drosophila melanogaster males with the agn ^ts3 mutation in the agnostic locus which contains CG1848 gene for LIMK1. The data obtained were compared with the results of our previous similar investigation on wild type CS males [1]. Flies were divided into 4 groups. The males of control groups were not subjected to heat shock. Other groups were comprised of males subjected to heat shock either at the beginning of larval development when the mushroom body neuroblasts are predominantly dividing (groups HS1), or at the prepupal stage when the brain central complex is developing (groups HS2), or at the imago stage 1 hour before the test (groups HS). All males were tested at the age of 5 days. Virgin and fertilized CS females were used as courtship objects. Comparison of control groups of the wild type (CS) and mutant strains has shown that the mutation agn ^ts3 does not affect the main parameters of courtship behavior (courtship latency, the rapidity of achieving copulation, courtship efficiency) but leads to: a decrease in sexual activity, an increase in duration of sound trains in the songs and to a slight increase in the rate and stability of singing pacemakers. When compared to wild type, the agn ^ts3 males are more resistant to HS given 1 hour before the test. After HS their courtship intensity does not decrease and the main parameters of their courtship behavior and communicative sound signals either do not change, or appear to be even better stabilized. The frequency of distorted sound pulses (an indicator of frequency of impairments in the activity pattern of neuro-motor circuits of the singing center) decreases, and the working rate and stability of pacemakers of the pulse and the sine songs increases. Therefore, although the sharply elevated LIMK1 and p-cofilin levels detected by immunofluorescent techniques in the cells of agn ^ts3 mutants can lead to dramatic defects in learning and memory [2] accompanied by a decline of motivation of males, they do not appreciably affect the neuro-motor coordination during singing. The higher resistance of characteristics of the mutant behavior and communicative sound signals to heat shock is in agreement with the fact that the extremely high LIMK1 and P-cofilin levels fall down to normal values after HS and both the learning acquisition and memory formation are restored.     

Mate guarding and male mate choice in the chameleon grasshopper <Emphasis Type="Italic">Kosciuscola tristis</Emphasis> (Orthoptera: Acrididae)

In the wild, male chameleon grasshoppers (Kosciuscola tristis) are frequently observed mounted on the back of females even when not in copula, and will fight off other usurping males. If this behaviour is mate guarding and reflects investment in male mate choice, then we expect males to preferably guard females based on reliable cues of quality. Cues for female quality likely include female size and egg development that together may indicate fecundity. We investigated male mate choice in the field expressed as mate-guarding preference, by comparing size and egg development in guarded and unguarded females. We found no difference between guarded and unguarded females in measures of fecundity or body size. The majority of females sampled did not contain any viable eggs. This finding suggests that male K. tristis indiscriminately guard females in a scramble mating system.     

Effects of male apparent length on female preference for absolute body size in <Emphasis Type="Italic">Xiphophorus helleri</Emphasis>

Female green swordtails (Xiphophorus helleri) prefer males of longer body and sword lengths, which may reflect perceptual constraints. Larger males project a larger image onto the female’s retina at a given viewing distance, which may elicit greater stimulation of the visual system and a stronger behavioral response. This study investigates the effects of male apparent size, as affected by proximity and sword length, on female preference for absolute body size in X. helleri. Females were presented pairs of dummy males in ten treatments organized into six categories: (1) the Actual Length Control where identically sized males were held at different viewing distances from the female; (2) the Sword Compensation Treatment in which viewing distance and apparent length (body + sword) were held constant, pairing sworded with swordless dummies of equal total length; (3) the Body Length Preference Treatment where male distance from the female was held constant, while varying body size; (4) the Apparent (Sword) Length Preference Treatments where male distance from the female was held constant, while varying sword length; (5) the Apparent Length Controls where body length, sword length, and viewing distances varied while holding apparent length constant; and (6) the Experimental Treatment in which the smaller-sworded male of the pair appeared larger. In each of the Actual Length Control, Body, and Sword Length Treatments, females preferred the closer, larger-bodied, and longer-sworded males, respectively. However, females exhibited preferences in neither any of the Apparent Size Control treatments nor any of the Sword Compensation Treatment. Additionally, females in the Experimental Treatment preferred the apparently longer-sworded dummy to the male whose sword was actually longer. These results suggest that female preference for male body and sword length is based on the male’s apparent rather than actual size.     

Effect of gut bacteria on fitness of the Chinese citrus fly, <Emphasis Type="Italic">Bactrocera minax</Emphasis> (Diptera: Tephritidae)

A likely symbiotic association between tephritid fruit flies and gut bacteria has been recognized since the beginning of the last century. However, direct evidence for a link between gut bacteria and fruit fly fitness is still limited or absent for many species. Similar to other tephritids, the gut of Bactrocera minax (Enderlein) (Diptera: Tephritidae) is known to contain bacteria throughout the life stage, but what, if any, impact these bacteria have on B. minax fitness is entirely unknown. In order to elucidate the effects of bacteria on the fitness of B. minax, resident bacteria were isolated from the adult gut using culture-dependent techniques. Adult fly diets were subsequently supplemented with three bacterial isolates (Klebsiella pneumonia, Citrobacter braakii and Pantoea dispersa), or bacteria were removed from flies by antibiotics treatment: untreated adults provided a control. Adult fitness parameters (male and female longevity, female fecundity, male copulation number) were measured for the two treatments and one control group. Results were complex depending on the fitness parameter measured and the bacterial species. Compared to the controls, antibiotic treated B. minax had significantly decreased fecundity, but male and female longevity was increased. When flies were fed diets supplemented with any of the three bacterial isolates, female fecundity was significantly enhanced. However, only Citrobacter braakii significantly increased male mating frequency than control males. The results show that gut bacteria directly influence fitness of B. minax, but impacts are dependent on the bacterial species and the fitness parameters measured.     

No Evidence of Intersexual Kin Recognition by Males of the Neotropical Paper Wasp <Emphasis Type="Italic">Polistes versicolor</Emphasis>

We investigated the effect of kinship on mate selection by males of the neotropical primitively eusocial paper wasp Polistes versicolor. By conducting short-time, paired, consecutive, dyadic encounters, in which a male was randomly introduced separately to a related and an unrelated female in a small glass arena, we were able to observe and measure the male sexual display. We found that male sexual interest toward a female was not affected by kinship, even after controlling the differences in female behavior. Despite our finding that males were as eager to harass females, regardless of kinship, previous evidence suggested that incestuous copulation (inbreeding) in natural populations of P. versicolor appeared to be avoided. We suggest that in the natural mating habitat of the native population that we studied, in which many colonies were able to produce males and reproductive females in a short time window, the probability of finding related sexual partners was low, hence, intersexual kin discrimination did not evolve.     

Dorsal Forewing White Spots of Male <Emphasis Type="Italic">Papilio polytes</Emphasis>(Lepidoptera: Papilionidae) not Maintained by Female Mate Choice

Palatable animals sometimes mimic the color patterns of noxious animals to gain protection from predators. This phenomenon, known as Batesian mimicry, is seen in many butterflies of the genus Papilio, and in some species the mimicry is limited to females. Although female-limited Batesian mimicry has been hypothesized to be caused by females preferring to mate with non-mimetic males, this hypothesis is rarely tested. In this study, we tested whether female mate choice is driving female-limited Batesian mimicry in Papilio polytes. Males have white spots on the dorsal forewings, which are absent in mimetic female sand in the toxic model, Pachliopta aristolochiae. Hence, we conducted mate choice experiments to examine whether these white spots are important to females. We offered females a choice of males with intact dorsal forewing white spots and males with artificially blackened dorsal forewings, resembling the model. Females did not show a preference for males with the white spots, suggesting that they are not being maintained by female mate choice. Future studies should investigate the presence of female mate choice on other parts of males’ wings to further understand the role of female mate choice, as well as explore other factors driving female-limited mimicry in these butterflies.     

Intersexual mimicry and flowering phenology facilitate pollination in a dioecious habitat specialist species, <Emphasis Type="Italic">Myristica fatua</Emphasis> (Myristicaceae)

Myristica fatua is a dioecious specialist species restricted to the endangered, freshwater Myristica swamp forests in the Western Ghats, India. Earlier studies have alluded to pollination by deception in members of the Myristica genus, and thus we examined the pollination ecology comprising floral biology, flower production, flower visitors, and reproductive success in M. fatua and inferred the potential strategies that could permit such deception in this habitat specialist tree. Male flowers provide pollen rewards for an extended period of time while female flowers are rewardless and both sexes are visited by generalist insects, mainly by honeybees and stingless bees. Bee visits were significantly more frequent and longer on male than on female flowers as bees collected pollen from male flowers. We found that flower production patterns create a preponderance of males compared to females in the swamp populations. Using a model of honeybee color vision, we found the distance between the color loci of male and female flowers and based on minimum visual angle subtended by these flowers, we suggest that the two floral sexes cannot be discriminated by bees. Bees are likely deceived by the perceptual similarity of rewardless female flowers to pollen-offering male flowers and pollination is the consequence of foraging errors made by pollinators that encounter largely male–rarely female flower mosaics as they forage among clump-distributed M. fatua trees in the swamp habitat.