The occurrence and function of the sphragis in the zerynthiine genera Zerynthia,Allancastria and Bhutanitis (Lepidoptera: Papilionoidea: Papilionidae)

In some butterfly species males attach a large external mating plug termed a sphragis to the female abdomen during mating. This is derived from male accessory secretions and covers the female ostium bursae and surrounding areas, thus preventing or delaying remating. Specimens of all 12 species of the genera Zerynthia, Allancastria and Bhutanitis (Lepidoptera: papilionidae), which form a natural clade within the Zerynthiini, were examined for presence or absence of a sphragis and their male and female genitalia were studied. In all three genera female genitalia lack a typical sinus vaginalis and the sterigma is modified to form an exposed, shiny, well-sclerotized genital plate, derived from the fusion and expansion of the lamellae ante- and postvaginales. The exposed ostium bursae is situated near the posterior end of the genital plate in Zerynthia, whereas in Allancastria and Bhutanitis it is near the anterior end. A crude irregularly formed sphragis was found at least facultatively in all species. The sphragides of Zerynthia and Bhutanitis were generally poorly developed, in most cases only partially covering the female genital plate. In Allancastria the sphragis mostly covered the genital plate entirely, and generally incorporated numerous long scales derived from the male’s 8th abdominal segment; scales were sometimes sparse or absent, probably due to depletion from repeated matings by males. In Zerynthia, males lacked the dense terminal abdominal tuft of elongated scales found in Allancastria, and their sphragis lacked scales. The sphragis of Bhutanitis thaidina incorporated scales from the male valves, whereas in the B. lidderdalii sphragis (and probably B. ludlowi) the scales derived from the male’s 8th abdominal segment. The role of the scales and possible reasons for the difference in the development of the sphragis among these genera are discussed.     

Breeding behaviour and mating success of Phyllomedusa rohdei (Anura,Hylidae) in south‐eastern Brazil

Observations on the courtship behaviour, mating behaviour and mating success of a leaf‐frog, Phyllomedusa rohdei, population were conducted in a temporary flooded site of the Atlantic Forest, south‐eastern Brazil. We did not find any influence of size and body weight on male mating success across three scales: seasonal, nightly chorus and local groups (males found within 1?m of a mated pair). In addition, no evidence was found that female Phyllomedusa rohdei exhibit overt mate choice. The formation of mated pairs in P. rohdei occurred because males moved toward the females, rather than reverse. The only variable that was significantly related to male mating success was the number of nights of participation in chorus activity. Besides calling tactics, males showed active searching behaviour. There was no evidence of size‐ and mass‐assortative mating, and a high percentage of fertilization at all size and mass ratios of males and females was observed. Clutches in this species are produced in the same manner found in others members of Phyllomedusa; a purse‐like nest is made over water by folding a leaf around the egg mass (embryos and fluid‐filled capsules). Finally, we supply a list of several studies that examined the relationship between size or mass ratios of males and females in amplexus and its influence on fertilization rate, and show that size‐assortative mating in anurans is inconsistent (present in some species but not in other).     

Studies on the life cycle of the cicadellid bug Empoasca dolichi Paoli,in Southern Nigeria

Summary

Studies carried out on the biology of E. dolichi Paoli showed that its eggs are palegreen to creamy white, elongate and often curved, and have a mean length and width of 0·84 mm and 0·16 mm respectively. The egg-incubation period averaged 8·3 days under uncontrolled insectary conditions of 21–32°C and 45–95% relative humidity, while the life-cycle from egg to adult averaged 17·5 days. In the insectary, mating in E. dolichi usually started between the third and sixth day after adult emergence. Over 56% of observed individuals mated on the 4th day. The observed diurnal mating period was mainly within 05·30–09·00 hours and 16·00–20·00 hours local time, with an average copulation period of 65·7 min. The mean pre-oviposition period of 3·0 days for 23 mated females was significantly shorter than 4·0 days for 21 unmated females. An average of 116 eggs and 96 eggs were laid by mated and unmated females respectively, but the fecundity rate (3 eggs/day of total life-span) of mated females was significantly higher than that (2 eggs/day) of unmated females. Unmated females lived longer (mean=49·7 days) than mated females with a mean of 38·3 days. The average life-span for mated and unmated males was 31·0 and 32·4 days respectively. Insectary and field observations revealed that females of E. dolichi preferentially lay their eggs into the stems, leaf-veins and leaf petioles of cowpea plants. Only mated females of E. dolichi laid viable eggs, and the observed male: female sex-ratio of their progeny in the insectary was 1:1·04.     

Copulatory courtship and morphology of genitalic coupling in seven Phyllophaga species (Coleoptera: Melolonthidae)

Males in six of seven species of Phyllophaga perform apparent courtship behaviour during copulation. As predicted by the hypothesis that such copulatory courtship is under sexual selection by female choice, behavioural details differ between species. The modified male abdominal sternites, which rub against the female's pygidium during copulation and whose morphology also differs between species, and the flattened, setose tarsi of males of one species probably function as contact courtship devices during copulatory courtship. Observations of events inside copulating beetles were made through the semi-transparent abdominal sternites of females and male genitalia in three species. They revealed cryptic genitalic thrusting behaviour of male genitalia within the female which occurred even when there was little or no external movement of the male genitalia. Dissections of pairs frozen in copula demonstrated that males transfer large quantities of material to females during copulation that is not physically related to sperm transfer. The type of material varied between species. In at least two species most of the transfer apparently occurred after sperm transfer. The possibility that some male genitalic structures function as holdfasts and/or stimulatory devices was suggested by their positions relative to female structures, and/or their movements within the female. Portions of male genitalia with strong bristles tended to lie against setose or more heavily sclerotized portions of the female reproductive duct. Some portions of female genitalia which vary between species seem ill-designed to exclude the genitalia of cross-specific males. The female reproductive tract is partially everted during copulation in one species.     

Mating behaviour and sexual selection in the African ball-rolling scarab Khepher platynotus (Bates) (Coleoptera: Scarabaeidae)

Mating behaviour of the ball-rolling dung beetle Kheper platynotus (Bates), adults of which are active in rainy seasons, was studied in southeastern Kenya. Male mating behaviour changed with the progression of the rainy season. In the early season when females made food balls, males attempted forced copulation with wandering and food-ball-making females on dung mounds. As the season progressed, females made brood balls, and males attempted to pair off with such females, as well as attempting forced copulation with wandering and food-ball-making females. A male who paired off with a brood-ball-making female cooperated with the female to make the ball. The male usually copulated after burying the ball (pre-copulatory mate guarding), but sometimes copulated during ball-making. In the latter case the male usually showed post-copulatory mate guarding until ball burial, but sometimes left soon after copulation. In the late season, when available females decreased because of maternal care in the underground nests, males started to make food balls to secure food resources for survival until the next rainy season. Male mate guarding involving ball-making, -rolling and -burying seemed to function to raise paternity confidence, but such assistance seemed to be less beneficial to females. Fights frequently occurred between guarding and intruding males on brood balls. Both residency and relative body sizes of fighters were important asymmetries influencing contest outcome. This seemed to result in not active but passive mate choice by brood-ball-making females.     

Long copulations and post-copulatory ‘escort’ behaviour in the locust leaf miner,Odontota dorsalis (Coleoptera: Chrysomelidae)

The adult behaviour of Odontota dorsalis on its host plant Robinia pseudo-acacia was studied by following marked individuals in nature. Copulations often lasted several hours, and were characteristically followed by long escort phases in which males remain mounted atop females. Pairing lasted as long as several days. Most pairs broke up around the time of day when most females were ovipositing. Females normally laid a small cluster of eggs once per day. Both sexes mated repeatedly, and females did not resist copulations.

Long pairings in O. dorsalis may have evolved because the encounter rate between the sexes is high and costs to the female low. It is suggested that escort behaviour will be found in many other species with similar ecology and in which males remain mounted on females while copulating.     

Antennae and the role of olfaction and contact stimulation in mate recognition by males of the pollinating fig wasp Ceratosolen gravelyi (Hymenoptera: Agaonidae)

Finding a mate is a crucial step in sexual reproduction. The mutualism between figs and their pollinators is exploited by many figwasps that develop within their galls and attain sexual maturity simultaneously. Male pollinators carry the specific burden of detecting and opening mating holes in galls containing conspecific females before mating with them. The mating behaviourand antennal sensilla of male pollinators associated with Ficus semicordata was investigated to understand mate recognition. Male Ceratosolen gravelyi located female-containing galls from a distance, but only attempted to chew a mating hole after antennal contact with the gall surface was made. They showed similar responses to females and body extracts but failed to respond to washed female bodies. This behaviour indicates that unidentified chemicals present on the body surfaces of females are sufficient to elicit olfactory attraction and tactile confirmation. Multiporous plate sensilla are candidates for olfactory reception from a distance while basiconic sensilla may be involved in contact chemoreceptor. All of the sensilla are highly localised, on the distal part of the terminal flagellomere, suggesting a response to selection for strong directionality in the complex odour environment of the fig in which messages produced by hundreds of females may confuse precise mate localisation.     

The scramble competition mating system of the sphecid wasp Palmodes praestans (Kohl)

Males of the wasp Palmodes praestans engage in non‐territorial patrolling behaviour within a scramble competition mating system, showing great fidelity to their wide‐ranging patrolling routes without attempting to monopolize the areas covered. Scramble competition appears adaptive given that mating opportunities are not spatially aggregated in this species, whose females build isolated, single‐celled nests. Because of the female distribution pattern, a male's reproductive success in P. praestans probably depends upon the ability to find highly scattered, unmated females. The females of some close relatives of this sphecid wasp form fairly dense nesting aggregations. As mating system theory predicts, the males of some (but not all) of these species focus their searching at these concentrated nesting/emergence sites where potential mates are clustered spatially.     

Mating behaviour and maternal care in the tropical savanna funnel-web spider Aglaoctenus lagotis Holmberg (Araneae: Lycosidae)

Despite their abundance and diversity, Neotropical spiders have been the subject of relatively few studies of behavioural ecology. The sexual behaviour and maternal care of the spider Aglaoctenus lagotis (Lycosidae) is described here. We performed experimental manipulations to test the hypothesis that males are attracted by female cues in the web. Two predictions were tested: (a) webs used by a virgin female will attract more males than webs used by an inseminated female; (b) the main cue in the web is chemical, so webs recently used by virgin females will attract more males than those that are not recently used. Additionally, we tested the hypothesis that maternal care increases offspring survivorship. Results showed that males are attracted by recently used webs (z = 0.0015; p = 0.037; z-test) and webs of virgin females (z = 0.041; p = 0.036, z-test). Male sexual behaviour presented three different categories: court, pre-mate and mate. After the mating, the females built a cocoon that was transported adhered to the spinnerets and held by the last pair of legs. After the hatching of the eggs, the young migrated to the dorsal region of the mother's body for 5 days. During maternal care the mother eliminates a yellowish drop from its oral cavity, which is collected by the young. During the development of the young until adulthood, the offspring that have their mother present in the first stages of life have greater survival compared with those without maternal care. Our results make it possible to perform comparative analyses between a South American Lycosidae species and other species distributed worldwide.     

XL.—A new Fossil form of Phasmatodea from Galkino (Turkestan), and on Mesozoic Phasmids in general

Insects of many groups form mating aggregations on hilltops. Males of some hilltopping species behave like certain classic lekking vertebrates, notably in their defense of territories that are devoid of resources useful to females but which are visited by females solely for mating. But even within a sample of 11 hilltopping insects found in the same habitat in central Arizona, there is considerable diversity with respect to resources transferred to mates by males, the degree to which males are aggregated and territorial, and the extent to which females can choose freely among hilltopping males. Thus ‘hilltopping’ is not a single mating system type; although some species conform closely to the traditional definition of a lek, others are very different. The ecological basis for the diversity in the behaviour of male hilltopping insects appears linked to differences in population density, which affect the costs of territoriality, and differences in the nature of female choice, which are little understood as yet.     

The genus Grammostola Simon 1892 (Araneae: Theraphosidae): a new species from western Argentina,new synonymy and distributional data

Explosive breeding is a common reproductive strategy, especially in anurans. In some cases of explosive breeding in anurans, intense aggressive interactions occur among males attempting to gain access to females, and for terrestrial species that mate in water, females may die by drowning during long mating struggles. Such occurrences are obviously detrimental to females, and may also cause a decrease in the reproductive success of males through depletion of energy without obtaining access to a live female. However, the males of the small Amazonian frog Rhinella proboscidea can promote the ejection of oocytes from the abdominal cavities of dead females and fertilize them. This behaviour can minimize losses to both partners during explosive reproduction events. The existence of such a “functional necrophile strategy” shows that there may be possible selection in favour of stronger and more persistent males in explosive breeders.     

Reproductive ecology of the glass frog Espadarana prosoblepon (Anura: Centrolenidae) in an urban forest of the Central Andes of Colombia

Population and reproductive behaviour aspects of the glass frog Espadarana prosoblepon are well known for populations from Central America, but similar information is lacking for populations from South America. We recorded the reproductive ecology of a population of the glass frog E. prosoblepon in the city of Armenia, Central Andes of Colombia. With weekly surveys with mark-recapture between December 2013 and November 2015, we monitored activity patterns and evaluated if the probability of males mating is associated to their body size or to chorus tenure (i.e. the number of nights a same individual is calling for attracting a mate). In addition, upon observing an amplexus, we recorded the behaviour of the individuals until oviposition and noted characteristics of egg clutch. We recorded 47 males and 32 females, nine amplexus events, eight ovipositions, and 21 egg clutches (35.4 ± 4.79 eggs per clutch, N = 29). Activity of males and females and the number of egg clutches were positively correlated to rainfall. Mating success of males was not associated to their body size, but it was positively associated to longer chorus tenure; clutches from large females had a higher number of eggs than clutches from small females. Oviposition occurred on average 3.76 ± 1.74 hours after we first observed the amplexus, occurring 93.1% of the time in fronds of Selaginella geniculata at an average height of 1.58 ± 0.44 m. After the oviposition, the male left the site, while the female remained between 30 and 45 min, partially covering the eggs. The reproductive behaviour of E. prosoblepon did not vary widely between a population in Costa Rica and our population in time to oviposition, and mating success of males associated to chorus tenure; however, in our study population we recorded a larger clutch size and the preference for S. geniculata fronds as oviposition substrate.     

“Riding” behaviour by males of Conops quadrifasciata (Diptera: Conopidae): Do females set up “riders” as targets for takeovers by larger males?

Males of Conops quadrifasciata “claim” females in late morning and then spend the rest of the day exposed as “riders” to attacks by other males. Male riders can respond to an attack by performing an abdomen‐lift which denies the attacker access to the female's genitalia. Attacking males can tailor their behaviour to suit the prevailing conditions. If a rider is engaged in a “repeat” copulation (as he does at regular intervals), the attacker “waits” beside the rider. If the rider is not in genitalic contact with the female, the attacker tries to mate with her immediately. It is proposed that females deliberately expose their riders to takeovers in order to gain the fittest mate.     

An imperfect imago? Post-mating loss of iridescent scales in Cheimas butterflies may change female from attractive to cryptic (Lepidoptera: Nymphalidae: Satyrinae)

Adults of both sexes of Cheimas opalinus (Staudinger), a Satyrinae butterfly occurring in the cloud forests of the Venezuelan Cordillera de Mérida, have simple wing colour patterns, dark brown dorsa marked by one conspicuous element, large greenish-blue patches in the discal area of the hind wing. It was observed that in young, freshly emerged females these patches are frequently absent. They fade away and eventually disappear during mating and nuptial flights when brushed out by lateral movements of the male’s hind wing put between folded female hind wings. The falling off of the scales is made easy because they do not adhere to the wing as firmly as brown background scales due to their particular folded extremities. We speculate that this process is related to the fitness of the females. Prior to mating, blue-green patches are advantageous because they attract the attention of the opposite sex and enhance the chances of successful mating. After mating they lose their sexual role. Females without the blue-green patches become cryptic. Thus, they are less apparent not only for the males, which helps them avoid sexual harassment, but also for potential bird predators. Additionally, the loss of blue reflecting scales may speed up the warming up of the abdomen and egg maturation. In the males, positive role of the patches (signalling), and negative (bird attraction), is balanced during their entire life span, and they fade gradually due to regular usage of wings scaling in flight. Our hypothesis is supported by morphological, optical, experimental and statistical analysis in which we used 509 individuals of both sexes. A similar ratio of young (with undamaged wing) females with or without blue patch was recorded, whereas in the males no individuals without any trace of blue patch were reported, and a correlation of age (wing damage) and gradual fading off of the patch was demonstrated. Such an adaptation involving an active change of the appearance of adults, i.e. colour patterns related to sexual selection, has not been reported previously in Lepidoptera.     

Expulsion and consumption of male ejaculates by promiscous female Euxesta eluta and Euxesta mazorca (Diptera: Ulidiidae)

While expulsion of male ejaculates by the females after copulation has been reported for various animal groups, expulsion followed by consumption of the expelled ejaculate is a rare behaviour outside spermatophylax-producing orthopterans. Among Diptera, this behaviour has been reported for a few species of Piophilidae, Empididae and Ulidiidae. Here we report on its occurrence among Euxesta eluta and Euxesta mazorca (Diptera: Ulidiidae). We also attempt to characterize the mating system of E. eluta in order to facilitate future hypothesis testing to understand the behavioural factors leading to the evolution of this peculiar behaviour. For this, courtship sequences, copulation duration, frequency of ejaculate expulsion and subsequent consumption, and latency to ejaculate expulsion for both E. eluta and E. mazorca were recorded. The time of sexual maturation, the time window of sexual receptivity during the day, and the mating frequency and variance in mating success for males and females (degree of polygamy) were determined for E. eluta. Both E. eluta and E. mazorca males engaged in elaborate courtship sequences involving visual and tactile displays before copulation. Females of both species almost invariably expelled and consumed ejaculates after copulation. Female E. eluta, required a 6- to 9-day period feeding on protein and sugar before becoming sexually receptive. Reproductive activity occurred continuously over the day with an early morning and late afternoon peak. Both males and females could mate multiply, with multiple partners over a 2-h observational period. Sexual network analysis revealed that some males and females had greater mating success than others. Ejaculate consumption appears to be widespread in the genus Euxesta. It is possible that females obtain nutrients from this behaviour while exerting some control over egg fertilization.     

Spawning behaviour of Engystomops pustulatus (Anura: Leptodactylidae)

Foam nests have evolved independently in several amphibian groups as an adaptive response to prevent predation and desiccation in dry environments. Nests are normally laid on ponds, or in underground galleries, humid forest leaf litter or terrestrial bromeliads. They are built when males or females beat a foam precursor associated with the egg masses extruded by the female. The spawning process requires the synchronic actions of the mating pair to obtain a hemispheric nest that protects the offspring. Herein, we describe the spawning behaviour of Engystomops pustulatus based on videos from 13 nesting couples from the lowlands of western Ecuador. Three variables were measured as indicators of male effort: duration of mixing events, duration of resting periods, and number of kicks per mixing event. We consider that not only male physical effort but also female behaviour influences nest structure. We suggest that nest building requires prolonged and intense physical activity by the male as well as the female’s steady position during spawning and female’s oviposition site selection. Nest building has two phases. In the first phase, the duration of resting periods, the duration of mixing events, and the number of kicks increase and are highly variable. During the second phase the three variables stabilise until the end. The volume of the nest increased mainly during the second phase. In four nesting events we observed kicking movements by the female. To our knowledge, this is the first time that female kicking has been observed in leptodactylid frogs. The function of this behaviour is unknown but our observations suggest that it may be triggered by insufficient male effort. Traditionally, female mate choice in Engystomops has been explained under models of indirect benefits exclusively. We argue that the prolonged male activity during nesting could influence female fitness directly. This will allow the operation of sexual selection via direct benefits.     

Courtship,copulation and oviposition in the chalcedon checkerspot,Euphydryas chalcedona (Lepidoptera: Nymphalidae)

In this report we describe some aspects of the reproductive biology of a population of the chalcedon checkerspot, Euphydryas chalcedona, in central Arizona. Successful courtship in this species lasts about a minute and has a form similar to that of other nymphalids. During copulation the male forms a spermatophore within the female's bursa copulatrix that represents about 7% of the male's body weight. After copulation females may remate and older females may have mated as many as three times. The spermatophore deflates as time since deposition passes. Females oviposit every other day on average and lay their eggs in clusters. These data confirm previous brief reports of courtship structure and spermatophore size for E. chalcedona. Questions are raised about the control of receptivity in this species and the function of the mating plug.     

Diurnal mating behaviour of a Nisitrus sp. cricket (Orthoptera: Gryllidae) from Sumatra

The sexual behaviour of a diurnally active cricket in the genus Nisitrus is described from observations made in its natural habitat. Courtship consists of both auditory and visual elements. The male transfers up to five spermatophores to an individual female during a single reproductive sequence lasting more than 3 h. Males guard females during the intervals between successive production of spermatophores. The visual signal-one or more hind-leg waves-given by the male to indicate to the female that she can eat the old spermatophore, and that a new one is now ready, appears to be unique among gryllids studied so far. Females are very co-operative, as they stand to receive and then eat as many as five spermatophores in relatively quick succession. Inter-male rivalry does not occur in the absence of females, but possible 'spoiling' behaviour may occasionally interrupt mating sequences.     

Interactions between the sexually deceptive orchid Spiculaea ciliata and its wasp pollinator Thynnoturneria sp. (Hymenoptera: Thynninae)

Males of the thynnine wasp Thynnoturneria sp. attempt to mate with female decoys in the flowers of the elbow orchid Spiculaea ciliata. Experimentally shifted orchids usually attract male wasps quickly, often within 2 minutes of presentation of the ‘bait’ orchids in appropriate habitat. Although the orchid effectively exploits the scramble competition mating system of the wasp, the insect is not totally at the mercy of the deceptive orchid. Fewer than half of all arriving males contact the column of the orchid flower, as required for orchid pollination. Moreover, the number of deceived visitors falls sharply over a short period and the number of wasp visitors does not rebound with the replacement of one bait orchid by another at that location. These observations suggest that patrolling wasp pollinators can discriminate to some extent between orchid decoys and female wasps, especially by learning to avoid particular locations that are associated with unrewarding flower decoys.     

The green June beetle (Cotinis nitida) (Coleoptera: Scarabaeidae): local variation in the beetle’s major avian predators and in the competition for mates

The green June beetle (Cotinis nitida) was observed in northern Virginia for a second and third flight season. The study yielded some results that were consistent with earlier findings, namely that mate-locating beetles were vulnerable to attack by avian predators and matings occurred primarily early in the flight season. Novel findings included the following: (1) blue jays (Cyanocitta cristata) were the major predators, not common grackles (Quiscalus quiscala), (2) some jays appeared to selectively prey on female beetles, (3) competition for mates among male beetles could be so intense that some males attempted to copulate with already mated (unreceptive) females, and (4) the sex ratio of the beetle population feeding on wild blackberries remained near equality late in the flight season despite the fact that the sex ratio of the population of beetles at the emergence site became highly female-biased over time.