The morphology of mouthparts,wings and genitalia of Paleozoic insect families Protohymenidae and Scytohymenidae reveals new details and supposed function

Megasecoptera is an extinct group of insects with specialized rostrum-like mouthparts, which is a synapomorphy shared with all members of the Late Paleozoic Palaeodictyopterida, and markedly slender wings that are unable to flex backwards. Here we describe the close up morphology of Protohymenidae and Scytohymenidae and uncover new aspects of the endoskeleton (tentorium) of the head, structure of the mouthparts with discernible proximal part of stylets controlled by muscles, surface of compound eyes that consist of a hexagonal pattern of large facets, structure and microstructures on the wings and reconstruct male and female external genitalia using ESEM and light stereomicroscopy. Furthermore, we describe Protohymen novokshonovi sp. n. based on an exceptionally well preserved fossil from the early Permian at Tshekarda in Russia, which shows crucial details, and the earliest species of Protohymenidae, Carbohymen testai gen. et sp. n. from a late Carboniferous siderite nodule at Mazon Creek in Illinois, USA. Our comparative study confirmed a set of structural and microstructural details on their wings, such as the composite anterior wing margin, development of an apical cell and the previously unknown external genitalia. Based on the results and comparison of homologous structures known primarily for extant relatives, such as mayflies and dragonflies, we outline for the first time the function of the mouthparts, in particular, the stylets, structure of the tentorium, vision provided by large hexagonal ommatidia and male copulatory structures bearing curved claspers for holding a female during copulation and penial lobes with seminal grooves.     

Paternity analyses in wild‐caught and laboratory‐reared Caribbean cricket females reveal the influence of mating environment on post‐copulatory sexual selection

Polyandry is ubiquitous in insects and provides the conditions necessary for male‐ and female‐driven forms of post‐copulatory sexual selection to arise. Populations of Amphiacusta sanctaecrucis exhibit significant divergence in portions of the male genitalia that are inserted directly into the female reproductive tract, suggesting that males may exercise some post‐copulatory control over fertilization success. We examine the potential for male–male and male–female post‐copulatory interactions to influence paternity in wild‐caught females of A. sanctaecrucis and contrast our findings with those obtained from females reared in a high‐density laboratory environment. We find that female A. sanctaecrucis exercise control by mating multiple times (females mount males), but that male–male post‐copulatory interactions may influence paternity success. Moreover, post‐copulatory interactions that affect reproductive success of males are not independent of mating environment: clutches of wild‐caught females exhibit higher sire diversity and lower paternity skew than clutches of laboratory‐reared females. There was no strong evidence for last male precedence in either case. Most attempts at disentangling the contributions of male–male and male–female interactions towards post‐copulatory sexual selection have been undertaken in a laboratory setting and may not capture the full context in which they take place – such as the relationship between premating and post‐mating interactions. Our results reinforce the importance of designing studies that can capture the multifaceted nature of sexual selection for elucidating the role of post‐copulatory sexual selection in driving the evolution of male and female reproductive traits, especially when different components (e.g. precopulatory and post‐copulatory interactions) do not exert independent effects on reproductive outcomes.     

External morphology of Xyleborus ferrugineus (Fabr.) (Coleoptera: Scolytidae). I. Head and prothorax of adult males and females

The overall body dimensions and external morphology of the head, head appendages, prothorax, and prothoracic legs of the adult haploid male and diploid female Xyleborus ferrugineus (Fabricius) were examined by scanning electron microscopy and light microscopy, and analyzed for variation. The female is significantly larger (1.26 X) than the male. The length/width ratio of both the pronotum and elytron, and the pronotum-length/elytron-length ratio are significantly different in the male and female. These findings indicate a distinct sexual dimorphism in general body form. There is significantly greater variability in pronotal length and width in the male than the female. There are sexual differences in setal patterns and setal size variabilities on the dorsal pronotum and the terminus of the antennae.     

Sexual selection in a moth: effect of symmetry on male mating success in the wild

Sexual selection is generally caused by female choice and male–malecompetition. In female choice process, female preference isfavored indirectly and/or directly by sexual selection. In indirectselection, females expressing the preference might gain indirectgenetic benefits. In direct selection, females expressing thepreference might gain direct benefits or avoid male-imposedcosts. The white-tailed zygaenid moth Elcysma westwoodii ismonandrous, and males often gather around a female to mate withher, suggesting a high opportunity for sexual selection on maletraits. We quantified phenotypic selection on male morphologyin this species in the field. The morphological characters analyzedincluded body weight, antenna length, forewing length, hindwing length, hind wing tail length, genital clasper length,and the fluctuating asymmetry (FA) of these bilateral traits.In E. westwoodii, selection favored males with more symmetricgenital claspers, as well as longer and more symmetrical hindwings and antennae. Negative correlations between FA and sizewere also detected in the clasper and the antenna. Our resultssuggest that FAs of male traits, in particular the genital clasper,may have indirect and direct influences on mating success. Duringa copulatory attempt, an E. westwoodii male will try to graspthe female's abdominal tip with his claspers but often failto do so because of the female's reluctance to mate. The femaleabdominal tips are smooth and strongly sclerotized and couldthus be difficult for males to grasp. We hypothesize that moresymmetrical male claspers are more efficient in overcoming femalereluctance.     

The male copulatory system of european pea crabs (crustacea,brachyura, pinnotheridae)

The male copulatory system of the European pinnotherid species Pinnotheres pisum, Pinnotheres pectunculi, and Nepinnotheres pinnotheres was investigated by gross morphology, scanning electron microscopy, histological methods, and confocal laser scanning microscopy. The brachyuran copulatory system is consistently formed by paired penes and two pairs of abdominal appendages, the gonopods, functioning in sperm transfer. In pinnotherids, the long first gonopods transfer the sperm mass into the female ducts. The first gonopod has the ejaculatory canal inside that opens both basally and distally. The second gonopod is solid, short, and conical. During copulation, the penis and the second gonopod are inserted into the basal lumen of the first gonopod. While the penis injects the sperm mass, the second gonopod functions in the transport of spermatozoa inside the ejaculatory canal toward its distal opening. The second gonopod is adapted for the sealing of the tubular system in the first gonopod by its specific shape and the ability to swell. Longitudinal cuticle foldings of the second gonopod hook into structures inside the first gonopod. The second gonopod can interact with the penis during copulation by a flexible flap separating the lumina in which the second gonopod and the penis are inserted. J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

The exoskeleton of the female genitalic region in Petrobiellus takunagae (Insecta: Archaeognatha): Insect-wide terminology,homologies, and functional interpretations

The exoskeleton of the female genitalic region (abdominal venters 7–9) in Petrobiellus takunagae (Machilidae-Petrobiellinae) is studied using light microscopy and SEM. Sclerites are distinguished from membrane by the degree of cuticular flexibility. However, the microsculpture of the cuticle is shown to be useful in characterising the heterogeneity of the cuticle and in detecting weak sclerotisations. The morphology of Petrobiellus is compared with that in Trigoniophthalmus alternatus (Machilidae-Machilinae) described previously. While venter 7 is similar, venters 8 and 9 show many differences in the presence/absence or fusion/separation of particular sclerites. This suggests female genitalic morphology to be a valuable character system for phylogenetic and taxonomic work in Archaeognatha. Comparison with other insect orders is aimed at detecting homologous structures and conditions. Important points are: (1) Petrobiellus has a sclerotised genital lobe posteriorly on venter 7, similar to Zygentoma and Dictyoptera; it bears the gonopore. (2) Petrobiellus has a posterior sclerite on venter 9 that is very similar to a sclerite of Odonata. (3) The morphology of the coxal lobes of venter 9 (gonoplacs) suggests their function as a sheath of the ovipositor. From female genitalic morphology we deduce the process of oviposition, describing an external egg transportation tract.     

Functional anatomy of male copulatory organs of Pomacea canaliculata (Caenogastropoda, Ampullariidae)

This study was aimed to investigate the functional morphology of copulation and sperm transfer in the invasive snail Pomacea canaliculata. Three-dimensional renderings of the male copulatory apparatus were made and showed elaborate systems for innervation and for hemolymph supply and drainage. A key component of the male copulatory apparatus is the penial sheath, which shows three specialized glands; the medial and distal glands may participate in adherence to the mantle cavity wall of the female during copulation. The outer gland has an epithelium composed of columnar cells with branched microvilli, mucous goblet cells and large granular secretory cells containing intragranular crystalloids, which produce an exocrine secretion during copulation. The interaction of male/female copulatory organs was studied in dissections of snap-frozen pairs. Sperm are left in the sperm pit, at the end of the pallial spermiduct. Afterwards, the muscular action of the penial bulb takes the sperm up to the vermiform penis, which slides from the penial pouch into the central groove of the penial sheath, and it later emerges through a T-shaped sulcus of this structure and enters the female vagina. Then, it climbs through the capsule duct, and its tip reaches the proximity of the seminal receptacle. A model of copulation and sperm transfer is presented on the basis of the new findings and on published literature.     

The function and evolution of male and female genitalia in Phyllophaga Harris scarab beetles (Coleoptera: Scarabaeidae)

Genitalia diversity in insects continues to fuel investigation of the function and evolution of these dynamic structures. Whereas most studies have focused on variation in male genitalia, an increasing number of studies on female genitalia have uncovered comparable diversity among females, but often at a much finer morphological scale. In this study, we analysed the function and evolution of male and female genitalia in Phyllophaga scarab beetles, a group in which both sexes exhibit genitalic diversity. To document the interaction between male and female structures during mating, we dissected flash‐frozen mating pairs from three Phyllophaga species and investigated fine‐scale morphology using SEM. We then reconstructed ancestral character states using a species tree inferred from mitochondrial and nuclear loci to elucidate and compare the evolutionary history of male and female genitalia. Our dissections revealed an interlocking mechanism of the female pubic process and male parameres that appears to improve the mechanical fit of the copulatory position. The comparative analyses, however, did not support coevolution of male and female structures and showed more erratic evolution of the female genitalia relative to males. By studying a group that exhibits obvious female genitalic diversity, we were able to demonstrate the relevance of female reproductive morphology in studies of male genital diversity.     

On the function of male genital claspers in <Emphasis Type="Italic">Stenomacra marginella</Emphasis> (Heteroptera: Largidae)

Male structures for clasping females during precopula interactions and mating (“claspers”) have evolved in many groups of arthropods. Several hypotheses regarding the function of claspers have been proposed. We describe how males of the true bug Stenomacra marginella (Heteroptera: Largidae) move their genital claspers during sexual interactions, and present the results of experiments in which we tested whether claspers are necessary to achieve intromission. When one and both claspers were partially amputated, the probability of successful intromission decreased from 62 to 0% and from 57 to 3%, respectively. Behavioral observations indicate that the claspers open the valves that cover the female genital opening. We consider the possibility that claspers in S. marginella may have multiple functions.     

Coevolution between Male and Female Genitalia in the Drosophila melanogaster Species Subgroup

In contrast to male genitalia that typically exhibit patterns of rapid and divergent evolution among internally fertilizing animals, female genitalia have been less well studied and are generally thought to evolve slowly among closely-related species. As a result, few cases of male-female genital coevolution have been documented. In Drosophila, female copulatory structures have been claimed to be mostly invariant compared to male structures. Here, we re-examined male and female genitalia in the nine species of the D. melanogaster subgroup. We describe several new species-specific female genital structures that appear to coevolve with male genital structures, and provide evidence that the coevolving structures contact each other during copulation. Several female structures might be defensive shields against apparently harmful male structures, such as cercal teeth, phallic hooks and spines. Evidence for male-female morphological coevolution in Drosophila has previously been shown at the post-copulatory level (e.g., sperm length and sperm storage organ size), and our results provide support for male-female coevolution at the copulatory level.     

Sexual Dimorphisms in the Dermal Denticles of the Lesser-Spotted Catshark,Scyliorhinus canicula (Linnaeus, 1758)

The dermal layers of several elasmobranch species have been shown to be sexually dimorphic. Generally, when this occurs the females have thicker dermal layers compared to those of males. This sexual dimorphism has been suggested to occur as a response to male biting during mating. Although male biting as a copulatory behaviour in Scyliorhinus canicula has been widely speculated to occur, only relatively recently has this behaviour been observed. Male S. canicula use their mouths to bite the female’s pectoral and caudal fins as part of their pre-copulatory behaviour and to grasp females during copulation. Previous work has shown that female S. canicula have a thicker epidermis compared to that of males. The structure of the dermal denticles in females may also differ from that of males in order to protect against male biting or to provide a greater degree of friction in order to allow the male more purchase. This study reveals that the length, width and density of the dermal denticles of mature male and female S. canicula are sexually dimorphic across the integument in areas where males have been observed to bite and wrap themselves around females (pectoral fin, area posterior to the pectoral fin, caudal fin, and pelvic girdle). No significant differences in the dermal denticle dimensions were found in other body areas examined (head, dorsal skin and caudal peduncle). Sexually dimorphic dermal denticles in mature S. canicula could be a response to male biting/wrapping as part of the copulatory process.     

Embryogenesis of the damselfly Euphaea yayeyamana Oguma (Insecta: Odonata: Euphaeidae), with special reference to the formation of their larval abdominal “gill‐like” appendages

The acquisition of wings in insects is the most significant subject in considering the diversification and adaptive radiation of insects, that is, the “macro‐evolution” of insects. In the discussion of the origin of insect wings, Palaeoptera has attracted particular attention in phylogenetic and evolutionary studies. In particular, Ephemeroptera have segmental gill‐structures on their abdominal segments during their nymphal stage, and these have been noted in discussions regarding their homology and/or serial homology between wings, gills and appendages. Although Odonata has received little attention in the course of these discussions, there are cases of segmental gill‐like structures on their abdomen in the two families, Euphaeidae and Polythoridae. Under such cirumstances, in this study, the embryological developmental process in Euphaea yayeyamana of Euphaeidae was observed, focusing on the formation process of the gill‐like structures. As a result, it was revealed that four of the seven pairs of gill‐like projection structures started their visible formation within the middle stages of embryonic development, and the remaining three pairs developed during the early stages of post‐embryogenesis. Some joint‐like structures existed in all of the gill‐like projections. It was revealed that muscle tissue was interposed within these protrusions and that all of the projections themselves fully articulated, and that the nervous system was extended into the protrusions. All of the gill‐like projections strongly suggested their homology with the cephalic and thoracic appendages, when we considered them with regard to their serial homology based on the topology of their formation position.     

Sexual dimorphism in calanoid copepods: morphology and function

Mate location and recognition are essentially asymmetrical processes in the reproductive biology of calanoid copepods with the active partner (the male) locating and catching the largely passive partner (the female). This behavioural asymmetry has led to the evolution of sexual dimorphism in copepods, playing many pivotal roles during the various successive phases of copulatory and post-copulatory behaviour. Sexually dimorphic appendages and structures are engaged in (1) mate recognition by the male; (2) capture of the female by the male; (3) transfer and attachment of a spermatophore to the female by the male; (4) removal of discharged spermatophore(s) by the female; and (5) fertilization and release of the eggs by the female. In many male calanoids, the antennulary chemosensory system is enhanced at the final moult and this enhancement appears to be strongly linked to their mate-locating role, i.e. detection of sex pheromones released by the female. It can be extreme in calanoids inhabiting oceanic waters, taking the form of a doubling in the number of aesthetascs on almost every segment, and is less expressed in forms residing in turbulent, neritic waters. Mate recognition is a process where chemoreception and mechanoreception presumably work in conjunction. The less elaborate male chemosensory system in the Centropagoidea is counterbalanced by females playing a more active role in generating hydromechanical cues. This is reflected in females in the shape of the posterior prosomal margin, the complexity of urosomal morphology and the size of the caudal setae. Visual mate recognition may be important in the Pontellidae, which typically show sexual dimorphism in eye design. The most distinctive sexual dimorphism is the atrophy of the mouthparts of non-feeding males, illustrating how copepod detection systems can be shifted to a new modality at the final moult. In the next phase, the male captures the female using the geniculate antennule and/or other appendages. Three types of antennulary geniculations are recognized, and their detailed morphology suggests that they have originated independently. Grasping efficiency can be enhanced by the development of supplemental hinges. The scanty data on capture mechanisms in males lacking geniculate antennules are reviewed. It is suggested that the loss of the antennulary geniculation in many non-centropagoidean calanoids has evolved in response to increasing predator pressure imposed on pairs in amplexus. Spermatophore transfer and placement are generally accomplished by the modified leg 5 of the male. In some males, leg 5 consists of both a chelate grasping leg and a spermatophore-transferring leg, whereas in others, only the latter is developed. Tufts of fine setules/spinules and/or sclerotized elements on the terminal portion of the leg are involved in the transfer and attachment of the spermatophore. The configuration of gonopores, copulatory pores and their connecting ducts in the female genital double-somite is diversified in the early calanoid offshoots such as Arietellidae and Metridinidae, whereas in more derived groups, it is constant and invariable, with paired gonopores and copulatory pores located beneath a single genital operculum. The absence of seminal receptacles in most Centropagoidea limits the female's ability to store sufficient sperm for multiple egg batches, suggesting that repeated mating is necessary for sustained egg production. Discharged spermatophores are usually removed by the female leg 5 and/or specialized elements on other legs. In Tortanus (Atortus) Ohtsuka, which has rudimentary fifth legs in the female and complex coupling devices in the male, a spermatophore supposedly remains on the female urosome, since eggs appear to be released from a ventral opening of the spermatophore. The type of sexual dimorphism is closely related to habitat and biology. Some hyperbenthic families never show multiplication of aesthetascs on the male antennule, whereas families of the open pelagic realm such as the Aetideidae always have non-feeding males exhibiting secondary multiplication of antennulary aesthetascs. The various aspects and diversity of calanoid sexual dimorphism are herein considered in an evolutionary context.     

Description of the male of Sclerocypris tuberculata (Methuen, 1910) (Crustacea,Ostracoda, Megalocypridinae)

The male of Sclerocypris tuberculata (Methuen), thus far unknown, is here described. Relying on the morphology of the copulatory appendages and of the prehensile palps, it appears that this taxon belongs to a separate species group, together with S. zelaznyi and perhaps also S. sarsi. There are some interesting sexual dimorphic characters in the valve morphology: males have shorter valves with a dorsal margin which runs nearly parallel to the ventral one (more elongated valves with sloping dorsal margin in females) and there is lobe-like projection of the valve margin on the ventro-caudal corner of the LV in females which is lacking in the male. Furthermore, the female genital region has a very aberrant morphology, and all specimens from the present collection possess the tuberculated and noded valves.     

Sexual coercion does not exclude luring behavior in the climbing camel-spider <Emphasis Type="Italic">Oltacola chacoensis</Emphasis> (Arachnida,Solifugae, Ammotrechidae)

Sexual coercion in the form of forced copulation has been used as a typical example to illustrate the conflict of interests between females and males. Among arthropods, forced copulation has been reported for some groups of insects and crustaceans, but not for arachnids. In the present work, we analyse and describe the behavioral patterns of mating behavior of the climbing camel-spider, Oltacola chacoensis, relating it to relevant morphological features, In this species, the male forcefully clasps the female’s genital region with his chelicerae and locks her fourth pair of legs with his pedipalps. In some cases, the cuticle of the female’s abdomen was damaged by this cheliceral clasping. In contrast to other camel-spiders, the female O. chacoensis never remained motionless during mating, but continuously shook her body, opening her chelicerae notably towards the male. Despite this coercive context, males performed copulatory courtship (tapping with pedipalps) and females showed an apparent cooperative behavior (they remained still during a short period of the sperm transfer phase). These results strengthen the idea that sexual coercion (in the form of forced copulation) and luring behavior (in the form of copulatory courtship) are not two mutually-exclusive male’s strategies during a single copulation.     

Uterine position and the activation of male sexual activity in testosterone propionate-treated female guinea pigs

Fetal female guinea pigs were delivered surgically and their position within a uterine horn relative to male fetuses was noted. When adult they were ovariectomized, injected daily with testosterone propionate, and examined for male copulatory behaviors in response to the presence of a receptive female. Females that developed contiguous to a caudal male were functionally similar to those that had been situated between two males; both exhibited enhanced levels of mounting relative to females that were located contiguous to no males (with and without the presence of a caudal male) and to females contiguous to a cephalic male. Therefore, both contiguity and positioning of the male fetus are necessary conditions for prenatal masculinization of the female guinea pig. Lastly, four animals that had resided singly in the uterus displayed the lowest levels of copulatory behaviors.     

Mating Behaviour in Laevicaudatan Clam Shrimp (Crustacea,Branchiopoda) and Functional Morphology of Male Claspers in a Phylogenetic Context: A Video-Based Analysis

Clam shrimps are freshwater branchiopod crustaceans which often present complicated breeding systems including asexual reproduction (parthenogenesis) and mixed mating systems (in androdioecious species both selfing and outcrossing occurs due to the co-presence of hermaphrodites and males). Reproductive patterns of Spinicaudata, which contains most clam shrimp species, have received much attention. Another group of clam shrimps, Laevicaudata, which holds a key position in branchiopod phylogeny, has practically not been studied. As a part of the mating process, males clasp to the carapace margin of the females with a pair (or two pairs) of anterior trunk limbs modified as claspers. Previous studies have shown that clasper morphology is important in a phylogenetic context, and that some parts of the claspers in Spinicaudata and Laevicaudata may have undergone a remarkable parallel evolution. Here we have used video microscopy to study aspects of the mating behaviour, egg extrusion, and fertilization in Lynceus brachyurus (Laevicaudata). It is shown that fertilization is likely to be external and that the peculiar tri-lobed lateral lamellae of female''s hind body assist in guiding the egg mass to the exopodal egg carriers where they are collected by their distal setation. The functional morphology of the male claspers was studied in detail by close-up video recordings. The movable “finger” of the clasper bends around the female''s carapace edge and serves to hold the female during mating. The larger palp grasps around the female carapace margin in a way very similar to the movable “finger”, possibly indirectly providing sensory input on the “finger” position. A brief comparative study of the claspers of a spinicaudatan clam shrimp showed both similarities and differences to the laevicaudatan claspers. The presence of two pairs of claspers in Spinicaudata seems to give males a better hold of the female which may play a role during extended mate guarding.     

Female mate preferences on high‐dimensional shape variation for male species recognition traits

Females in many animal species must discriminate between conspecific and heterospecific males when choosing mates. Such mating preferences that discriminate against heterospecifics may inadvertently also affect the mating success of conspecific males, particularly those with more extreme phenotypes. From this expectation, we hypothesized that female mate choice should cause Enallagma females (Odonata: Coenagrionidae) to discriminate against conspecific males with more extreme phenotypes of the claspers males use to grasp females while mating – the main feature of species mate recognition in these species. To test this, we compared cerci sizes and shapes between males that were captured while mating with females to males that were captured at the same time but not mating in three Enallagma species. In contrast to our hypothesis, we found only one of forty comparisons of shape variation that was consistent with females discriminating against males with more extreme cerci shapes. Instead, differences in cerci shape between mating and single males suggested that females displayed directional preferences on 1–4 aspects of cerci shape in two of the species in our samples. These results suggest that whereas some directional biases in mating based on cerci shape occur, the intraspecific phenotypic variation in male cerci size and shape is likely not large enough for females to express any significant incidental discrimination among conspecifics with more extreme shapes.