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.     

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.     

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.     

The function of the sphragis in Cressida cressida (Fab.) (Lepidoptera,Papilionidae): a visual deterrent to copulation attempts

During mating, males of Cressida cressida (Lepidoptera, Papilionidae) form a large external mating plug or sphragis that covers the female's copulatory opening and physically prevents remating. The sphragis has lateral and distal projections that make it an obvious structure. We tested experimentally the hypothesis that the sphragis acts as a signal of female mating status. Males pursue mated females with an intact sphragis only briefly and rarely make physical contact with them. When the sphragis of a mated female is removed or reduced in size, males are significantly more likely to physically contact the female and initiate the aerial takedown that preceeds a copulation attempt. These results suggest that the sphragis deters male sexual interest at a distance and thus functions as a signal of female mating status. The discussion focuses on the fitness consequences for females and their mates of the signal function of the sphragis.     

Palpal loss,single palp copulation and obligatory mate consumption in Tidarren cuneolatum (Tullgren, 1910) (Araneae,Theridiidae)

Males of Tidarren cuneolatum (Tullgren, 1910) amputate one of their palps a few hours after the penultimate moult, like T. sisyphoides (Walckenaer, 1841) from the USA (Branch, 1942). Hence adult males, which are minute, have only one palp, either left or right randomly. This palpal organ is not oversized, when compared with other small spiders. During courtship females are unusually active, signalling receptivity by continuous twanging with legs II. Males construct a mating thread. Copulation involves one insertion, which lasts ca 4 min. Thus, only one receptaculum is inseminated during copulation. With the advance of insertion the male's prosoma becomes shrunken. Copulation regularly ends in mate consumption. In copulation with a virgin female the palp was inserted contralaterally. Females taken in the field had both receptacula filled with sperm and therefore were polyandrous. Re-mating was also observed in the laboratory. Remarkably the second male performed an ipsilateral insertion, if it possessed the same palp as the previous male. Probably the virgin receptaculum was recognized. Postembryonic development is rapid in males, which moult three times and mature ca 41 days after hatching from the cocoon. Females need four or five moults and ca 69 days to reach maturity and then survive ca 2-4 months.     

“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.     

Sexual dimorphism in chelicerae,forelegs and palpal traits in two burrowing wolf spiders (Araneae: Lycosidae) with sex-role reversal

Allocosa alticeps and Allocosa brasiliensis are two sex-role-reversed spiders. Females locate males and initiate courtship. Copulation occurs inside male burrows and after copulation the males deliver their burrows to the females. We explored the occurrence of differences in morphological traits related to sex roles in both Allocosa species and compared the results with two non-burrowing wolf spiders with typical sex roles. We measured the length of the foreleg's tibia-patella and the chelicerae. Scanning electron micrographs of palpal tarsus distal sections were taken. Males showed higher values than females in all the traits considered. Adult and penultimate males lacked true claws compared with juveniles and females, but showed modified spines. The palpal organ was more proximal to the tibia compared with the non-burrowing lycosids considered in this study. Spines and palpal organ location could be associated with more effective digging. We discuss how natural and sexual selection could interact on morphology in this scenario.     

Three new species of Pseudohaliotrema Yamaguti, 1953 (Monogenea: Ancyrocephalidae) from Siganus species (Siganidae) and the description of a mechanism for cross-insemination

Three new species of Pseudohaliotrema Yamaguti, 1953 were collected from two Siganus species caught off Singapore. They are Pseudohaliotrema virgata n. sp. and P. molnari n. sp. from Siganus virgatus and P. falcata n. sp. from S. guttatus. They differ in the shape and size of both the body and haptoral sclerites and in the morphology of the male organ (modified copulatory organ). Pseudohaliotrema species from siganids are characterized by having a large prominent vaginal pouch, an eversible spermatophore transfer tube associated with the vaginal pouch, ovoid spermatophores (in mature worms), a male organ adapted for spermatophore formation, a large prostatic reservoir, and a testis and ovary arranged side by side. Pseudohaliotrema plectocirra Paperna, 1972 is re-assigned as Tetrancistrum plectocirra (Paperna, 1972) n. comb. Pseudohaliotrema species from non-siganid fish do not possess these characteristics and belong elsewhere. The generic diagnosis of Pseudohaliotrema is amended. Pseudohaliotrema species use a spermatophore for sperm transfer, which is documented in detail for P. virgata n. sp. The same reproductive strategy is used by P. molnari n. sp. and P. falcata n. sp., as indicated by the presence of spermatophores on the spermatophore transfer tube and inside the vaginal pouch. The presence of individuals exchanging spermatophores indicates that P. sphincteroporus Yamaguti, 1953 and P. sigani Yamaguti, 1953 also employ the same mechanism. The spermatophore, assembled in the sclerotized male organ and stored in the spermatophore chamber, is retrieved by the eversible spermatophore transfer tube (housed within the vaginal pouch) and drawn into the vaginal pouch as the spermatophore transfer tube invaginates. Within the vaginal pouch, the wall of the spermatophore disintegrates releasing spermatozoa.     

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 in two closely related sympatric species of Philonthus (Coleoptera: Staphylinidae)

Summary

The intra-specific mating behaviour of two closely related sympatric staphylinid beetles, Philonthus cephalotes (Grav.) and P. sordidus (Grav.) is described in detail. Mating is essentially similar in the two species, although differing in the duration of copulation (1·7 min in cephalotes, 44·6 min in sordidus), and consists of four phases (A)-(D); (A) pre-copulatory phase, (1) examination of female abdominal apex by male, (2) examination of male abdominal apex by female, (3) prolonged examination of female by male, (4) mounting by male (5) extrusion of male genitalia; (B) copulatory phase; (C) terminal phase; and (D) post-copulatory phase.

An attempt to test the role of species specific patterns of peg setae, occurring on the genitalia of males, as a tactile signalling system was made by observation of inter-specific interactions between males and females of the two species. Under the trial conditions inter-specific intromission did not occur and behaviour was predominantly aggressive. Evidence suggests that signals during the early stages of an encounter between the sexes of different species are generally sufficient to deter further courtship. In two instances courtship reached a more advanced stage and here the patterns of peg setae on the male genitalia may have come into play, operating as a signalling device.     

The structure and function of the adhesive organs on the antennae of male Harpocera thoracica (Fallen) (Miridae; Hemiptera)

Summary

S.E.M. micrographs of unusual setae on the male antennae, and of the female pronotum in Harpocera thoracica (Fallen) (Miridae; Hemiptera) are presented. The adhesive function and positioning of the male antennae on the female's pronotum during pairing is discussed.     

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.     

Antennation and sexual performance of male digger wasps Sphex ingens Smith (Hymenoptera: Sphecidae)

Antennation can play several roles in hymenopteran copulation, mainly in wasps. The digger wasp Sphex ingens Smith has a very peculiar sexual behaviour, in which forced copulation is a striking element. However, communication through antennation during the pre-copulation and copulation phases and the relationship between the sexual performance of males and the final result of copulation require further clarification. The sexual behaviour of wild populations of S. ingens was filmed during the breeding season in a site between the beaches Meros and Aventureiro, Ilha Grande, southeastern Brazil. We assessed antennation behaviour, courtship duration, sexual performance, and genital clasping. Only successful males had the genital clasping assessed. Evidence found so far points out that the successful males showed better sexual performance – that is, they antennated more in a shorter courtship duration. Also, genital clasping in successful males did not depend on antennation, courtship duration, or sexual performance. Our results shed light on some aspects of the sexual behaviour of wasps, as they explain part of the sexual selection mechanisms adopted by the species.     

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.     

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.     

External morphology of Coptophylla caroliniani and Aceria mississippiensis spermatophores (Prostigmata: Eriophyidae)

The spermatophores of Coptophylla caroliniani and Aceria mississippiensis consist of a base which is glued to the substrate by an adhesive substance, a stalk, and an expanded head with a sperm packet resting on a central projection. The C. caroliniani spermatophore has a twisted stalk whereas the A. mississippiensis spermatophore has a straight stalk that is twisted near the junction of the head and stalk. A fan-shaped head is characteristic of a C. caroliniani spermatophore whereas A. mississippiensis has a shield-shaped head. The central projection which attaches the sperm packet to the head is oblong in C. caroliniani and cylindrical in A. mississippiensis.     

Intersex in the gonochoristic crustacean Tanais dulongii (Audouin, 1826) (Peracarida: Tanaidacea: Tanaididae): a comparison of external reproductive characteristics

Tanais dulongii (Audouin, 1826) is a small benthic crustacean with separate sexes and a marked sexual dimorphism characterized by a cephalothorax narrowed anteriorly, larger chelae (claws) and a pair of tiny genital cones in males, and ovisacs that form chambers for carrying embryos in females. Recent studies have described the existence of intersexual individuals with a cephalothorax similar to males, large chelae and ovisacs, but, due to their small size, it has not been possible to confirm the presence of genital cones under normal light microscopy. The present study analyses and compares external reproductive structures under scanning electron microscopy, of male and female T. dulongii, to corroborate the presence of both ovisacs and genital cones in potentially intersex individuals. The possible causes, such as hermaphroditism, environmental sex determination, parasitism, genetic aberration or endocrine-disrupting pollutants, are discussed.     

Are there costs to extended larval transport in the Trinidadian stream frog,Mannophryne trinitatis (Dendrobatidae)?

Previous work has shown that male Mannophryne trinitatis (Dendrobatidae) carry their larvae on their backs for up to 4 days in search of a predator‐free pool in which to deposit them. The experiments reported here investigated whether costs to the larvae or to the adults limit transport duration. We simulated transport durations of 0, 4, 8, and 12 days for larvae, but found no deterioration in terms of ability to grow to metamorphosis; indeed, 12‐day larvae grew better than all the others. After 8 days of simulated transport, larvae had used up all their yolk reserves and begun to lose dry weight. Larvae on wet substrates gained wet weight and length but on drier substrates merely maintained weight, suggesting that dehydration could be a problem on the male's back. In a trial of locomotor performance (mean jump length; number of jumps to traverse a runway), females performed best with calling males not significantly different from transporting males, despite an average larval load equivalent to 15–20% of the frog's mass. Assessment of gut contents showed that females foraged more than males, but that transporting males foraged as much as did calling males. We found no differences between the three classes of adult frogs in fat body weight.     

Two new genera of the family Pilumnidae (Crustacea: Decapoda: Brachyura) from Queensland,Australia

Two new genera of Pilumnidae each with a single new species are described from southeastern Queensland waters. Priapipilumnus is defined by the greatly swollen base and simple blunt apex of the male first pleopod and the concomitant posteriorly protruding, bilobed, third abdominal segment. Takedana is separable by the very short spinous anterolateral margins, the swollen branchial regions, the moderately short walking legs, and the strongly spined chelae. Actumnus pugilator is reassigned to Bathypilumnus Ng and Tan, 1984.     

Description of a new species of Asymmetrione (Isopoda: Bopyridae: Pseudioninae) infesting the hermit crab Paguristes tomentosus (Anomura: Diogenidae) from Peru,with a key to species and a review of southeastern Pacific bopyrids

A new species of bopyrid isopod is described from a host hermit crab Paguristes tomentosus collected from Pisco, Peru and assigned to the pseudionine isopod genus Asymmetrione Codreanu, Codreanu and Pike, 1965. This is the second species in the genus with less than 20° asymmetry in females, but agrees with all other species in the genus in having the characteristic “socket” on the propodus of the female's pereopods, and in all characters of the males. A key is provided to the 10 species now in Asymmetrione. New records of an Anathelges sp. from Pagurus villosus collected in central Chile are given but additional material is required to determine whether these specimens represent the eastern Pacific A. thompsoni or the western Atlantic A. hyptius. A review of all the bopyrid species known from the western South American coast, with remarks on their taxonomy and biology, is provided.