Interspecific variation in ovipositor morphology among Manaosbiid and Nomoclastid Harvestmen (Arachnida,Opiliones, Laniatores)

The external anatomy of the ovipositor has generally been overlooked as a source of informative characters in systematic studies of laniatorean harvestmen. In this study, we used scanning electron microscopy to examine the ovipositors of nine species representing the families Manaosbiidae (five species) and Nomoclastidae (four species). Similar to the ovipositor morphology of many gonyleptoidean families, the distal tips of the ovipositors of these harvestmen have four external lobes, with the margins most commonly adorned with 10 large peripheral setae. In manaosbiid and nomoclastid species, these peripheral setae have undivided bases, striated shafts and undivided distal tips. There are typically three setae on each anterior lobe and two setae on each posterior lobe. The medial setae on both anterior and posterior lobes insert into sockets that are slightly more dorsal. We observed small, surface denticles, usually associated with a pore, on the external surface of the lobes. There was interspecific and intraspecific variation in the number and shape of these surface denticles. The association of pores with denticles on the ovipositor appears to be a feature common to not only both families but is also a trait that has not been observed on ovipositors in other families of laniatorean harvestmen.     

Interspecific morphological variation in the wood-feeding cockroach, Cryptocercus (Dictyoptera: Cryptocercidae)

Appalachian species of Cryptocercus (Dictyoptera: Cryptocercidae) display considerable genetic variation, but little morphological variation has been reported. We employed light and scanning electron microscopy to investigate variation in male and female reproductive structures among four species of Cryptocercus. Our results indicated consistent, species-specific differences exist in the genitalia of the four species. Males exhibit moderate interspecific differences in the shape of the subgenital plate and third left phallomere hook. Females display consistent interspecific differences in the size of setae on the basivalvula and great differences in size, number, and pattern of setae on the base of the second valves. Together, the interspecific variation in female reproductive structures can be used to identify each of the four Cryptocercus species that occur in the Appalachian Mountains.     

Intraspecific and interspecific variation of female genitalia in two species of watersnake

The morphological differences in female genitalia within and between species are little studied and poorly understood, yet understanding patterns of variation in female genitalia can provide insights into mechanisms of genital evolution. The present study aimed to explore the patterns of intraspecific and interspecific variation in female genitalia in two sister taxa of watersnake (Nerodia sipedon and Nerodia fasciata) that have similar genital shape. We used a geometric morphometric (GM) approach to study variation in shape of the vagina between and within two sister species. We examined genital shape in female watersnakes ranging from small, sexually immature females to large reproductive females that had recently given birth. We found that shape variation of genitalia is strongly correlated with body size, where larger but not smaller females have a bifurcation in the vagina. However, we also found significant shape variation in the structure of the vagina between the two species, where N. fasciata has narrower genitalia with more prominent bifurcation, whereas N. sipedon has wider genitalia with less marked bifurcation. Using GM allowed us to detect significant differences in genital shape that were not apparent upon visual examination alone, suggesting that shape variation in female genitalia may be greater than previously assumed. Additional study of morphological differences in male reproductive organs for these species would help to determine whether there has been genital co‐evolution, and potentially mechanical reproductive isolation, in these two closely‐related and occasionally sympatric species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 183–191.     

The coevolution of male and female genitalia in a mammal: A quantitative genetic insight

Male genitalia are among the most phenotypically diverse morphological traits, and sexual selection is widely accepted as being responsible for their evolutionary divergence. Studies of house mice suggest that the shape of the baculum (penis bone) affects male reproductive fitness and experimentally imposed postmating sexual selection has been shown to drive divergence in baculum shape across generations. Much less is known of the morphology of female genitalia and its coevolution with male genitalia. In light of this, we used a paternal half-sibling design to explore patterns of additive genetic variation and covariation underlying baculum shape and female vaginal tract size in house mice (Mus musculus domesticus). We applied a landmark-based morphometrics approach to measure baculum size and shape in males and the length of the vaginal tract and width of the cervix in females. Our results reveal significant additive genetic variation in house mouse baculum morphology and cervix width, as well as evidence for genetic covariation between male and female genital measures. Our data thereby provide novel insight into the potential for the coevolutionary divergence of male and female genital traits in a mammal.     

No evidence for external genital morphology affecting cryptic female choice and reproductive isolation in Drosophila

Genitalia are one of the most rapidly diverging morphological features in animals. The evolution of genital morphology is proposed to be driven by sexual selection via cryptic female choice, whereby a female selectively uptakes and uses a particular male's sperm on the basis of male genital morphology. The resulting shifts in genital morphology within a species can lead to divergence in genitalia between species, and consequently to reproductive isolation and speciation. Although this conceptual framework is supported by correlative data, there is little direct empirical evidence. Here, we used a microdissection laser to alter the morphology of the external male genitalia in Drosophila, a widely used genetic model for both genital shape and cryptic female choice. We evaluate the effect of precision alterations to lobe morphology on both interspecific and intraspecific mating, and demonstrate experimentally that the male genital lobes do not affect copulation duration or cryptic female choice, contrary to long‐standing assumptions regarding the role of the lobes in this model system. Rather, we demonstrate that the lobes are essential for copulation to occur. Moreover, slight alterations to the lobes significantly reduced copulatory success only in competitive environments, identifying precopulatory sexual selection as a potential contributing force behind genital diversification.     

Multiple sexual selection pressures drive the rapid evolution of complex morphology in a male secondary genital structure

The genitalia of internally fertilizing taxa represent a striking example of rapid morphological evolution. Although sexual selection can shape variation in genital morphology, it has been difficult to test whether multiple sexual selection pressures combine to drive the rapid evolution of individual genital structures. Here, we test the hypothesis that both pre‐ and postcopulatory sexual selection can act in concert to shape complex structural variation in secondary genital morphology. We genetically modified the size and shape of the posterior lobes of Drosophila melanogaster males and tested the consequences of morphological variation on several reproductive measures. We found that the posterior lobes are necessary for genital coupling and that they are also the targets of multiple postcopulatory processes that shape quantitative variation in morphology, even though these structures make no direct contact with the external female genitalia or internal reproductive organs during mating. We also found that males with smaller and less structurally complex posterior lobes suffer substantial fitness costs in competitive fertilization experiments. Our results show that sexual selection mechanisms can combine to shape the morphology of a single genital structure and that the posterior lobes of D. melanogaster are the targets of multiple postcopulatory selection pressures.     

Interspecific variation in the microanatomy of cosmetid harvestmen (Arachnida,Opiliones, Laniatores)

Scanning electron microscopy (SEM) is a useful tool for identifying interspecific variation in often overlooked structures that may represent useful sources for informative phylogenetic characters. In this study, we used SEM to compare the morphology of 12 cosmetid species from Central America, the Caribbean, and North America including multiple species for the genera Cynorta, Erginulus, and Paecilaema. To determine if microanatomical structures were unique to the cosmetid taxa under examination, we investigated the microanatomical structures of six additional species of gonyleptoidean harvestmen representing the families Agoristenidae, Cranaidae, Gonyleptidae, Manaosbiidae, and Stygnidae. Our results indicate that the shape of the ocularium (narrow, intermediate, or broad) did not vary within cosmetid genera, whereas the morphology of the rough pit glands on the eye mound varied considerably between species. Each cosmetid species had 10–20 rough pit glands on the ocularium whereas only the eye mounds of Avima intermedia (Agoristenidae) and Glysterus sp. (Gonyleptidae) had similar structures. With regards to the surface texture of the dorsal scutum, cosmetid harvestmen exhibited a rivulose‐microgranulate morphology (6 species), a microtuberculate‐rivulose‐microrgranulate morphology (4 species), or a microgranulate morphology (2 species). In contrast, each of the gonyleptoidean species exhibited a microgranulate pattern, with the exception of Stygnoplus clavotibialis, which had a rivulose‐microgranulate surface texture. For cosmetid harvestmen, we observed considerable interspecific variation in the shape and number of teeth on the fixed and moveable fingers of the male chelicerae. Similarly, we also observed interspecific variation in the distribution and shape of tubercles on the ventral and dorsal surfaces of the femur of the pedipalp. Overall, our results indicate that there are several microanatomical structures associated with the ocularium, dorsal scutum, male chelicera, and pedipalp that could represent informative phylogenetic characters in future taxonomic studies of cosmetid harvestmen. J. Morphol. 275:1386–1405, 2014. © 2014 Wiley Periodicals, Inc.     

COPULATORY BEHAVIOR,GENITAL MORPHOLOGY,AND MALE FERTILIZATION SUCCESS IN WATER STRIDERS

Recent theoretical and empirical interest in postmating processes have generated a need for increasing our understanding of the sources of variance in fertilization success among males. Of particular importance is whether such postmating sexual selection merely reinforces the effects of premating sexual selection or whether other types of male traits are involved. In the current study, we document large intraspecific variation in last male sperm precedence in the water strider Gerris lateralis. Male relative paternity success was repeatable across replicate females, showing that males differ consistently in their ability to achieve fertilizations. By analyzing shape variation in male genital morphology, we were able to demonstrate that the shape of male intromittent genitalia was related to relative paternity success. This is the first direct experimental support for the suggestion that male genitalia evolve by postmating sexual selection. A detailed analysis revealed that different components of male genitalia had different effects, some affecting male ability to achieve sperm precedence and others affecting male ability to avoid sperm precedence by subsequent males. Further, the effects of the shape of the male genitalia on paternity success was in part dependent on female morphology, suggesting that selection on male genitalia will depend on the frequency distribution of female phenotypes. We failed to find any effects of other morphological traits, such as male body size or the degree of asymmetry in leg length, on fertilization success. Although males differed consistently in their copulatory behavior, copulation duration was the only behavioral trait that had any significant effect on paternity.     

Leg injuries and wound repair among cosmetid harvestmen (Arachnida,Opiliones, Laniatores)

Previous studies of leg injuries in harvestmen have focused on the fitness consequences for individuals that use autospasy (voluntary detachment of the leg) as a secondary defense mechanism. Leg damage among non‐autotomizing species of laniatorean harvestmen has not been investigated. Under laboratory conditions, we damaged femur IV of Cynorta marginalis and observed with scanning electron microscopy (SEM) the changes in these wounds over ten days. We also used SEM to examine leg damage from individuals of three species of cosmetid harvestmen that were collected in the field. On the basis of changes in the external surface of the hemolymph coagulum, we classified these wounds as fresh (coagulum forming), recent (coagulum with smooth surface), older (coagulum is scale‐like with visible cell fragments), and fully healed (scale replaced by new cuticle growth on the terminal stump). Our observations indicate that wound healing in harvestmen occurs in a manner comparable to that of other chelicerates. Leg injuries exhibited interspecific variation with respect to the overall frequency of leg wounds and the specific legs that were most commonly damaged. In addition, we measured walking and climbing speeds of adult C . marginalis and found that individuals with fresh injuries (lab‐induced) to femur IV walked at speeds significantly slower than uninjured adults or individuals collected from the field that had fully healed wounds to a single leg. J. Morphol. 278:73–88, 2017. ©© 2016 Wiley Periodicals,Inc.     

Phylogenetic and taxonomic problems in freshwater Oligochaeta with special emphasis on chitinous structures in Tubificinae

Setae and chitinous penis sheaths are the main characters used to distinguish genera and species of the subfamily Tubificinae. Genital setae and penis sheaths are of functional importance to facilitate copulation. Similar structures in different genera may be homologous or products of parallel evolution (homoiologous). Form and dimensions of the penis sheaths of many tubificine species are very variable. Transspecific overlap of quantitative species characters can make the determination of specimens in the genus Limnodrilus difficult. The configuration of the distal ends of the penis sheaths is an important character to distinguish Limnodrilus species. The definition of the intraspecific variability of this morphological character is problematic.     

Differences in genitalia structure and function between subfamilies of longhorn beetles (Coleoptera: Cerambycidae)

Beetle genitalia are usually described only for taxonomic purposes without considering the possible function of structures. Exceptions are sporadic detailed studies on single species. We studied genital structures in the subfamilies of Cerambycidae and outlined assumptions on the function of these structures and the implications for the phylogeny of the Cerambycidae. We found that male genitalia in particular are taxon-specific on a higher taxonomic level; e.g., the parameres are widely variable in Cerambycinae, while in most Lamiinae species they appear relatively uniform and differ from those of the Cerambycinae. Internal sac structures are very different among the various subfamilies. Small backwards-pointing spines are the most common armature of the internal sac. The female genitalia are less variable, although ovipositor morphology may differ among subfamilies. In most species, the connection between the mates during copulation is achieved by the long internal sac and the ovipositor only, whereas the median lobe and parameres are in contact with the female abdomen only at the beginning of copulation. Cerambycinae and Lepturinae have a basal swelling of the endophallus to prevent it from sliding back into the male abdomen during copulation. The long internal sac functions in connecting the mates and guaranteeing the sperm transfer.     

Arhopalus ferus (Coleoptera: Cerambycidae): structure and function of the female reproductive system

Abstract

Arhopalus ferus (Mulsant) female pupae and adults were dissected, and both internal and external genitalia are described. The external genitalia involve segments 8 and 9. Segment 9 and its appendages are modified into a long ovipositor which telescopes with segment 8 into segment 7 when retracted. The internal genitalia include ovaries, bursa copulatrix, spermatheca, spermathecal gland, vagina, and accessory glands. The terminology applied to coleopteran genital anatomy is critically discussed.     

Intraspecific evidence from guppies for correlated patterns of male and female genital trait diversification

The role of sexual selection in fuelling genital evolution is becoming increasingly apparent from comparative studies revealing interspecific divergence in male genitalia and evolutionary associations between male and female genital traits. Despite this, we know little about intraspecific variance in male genital morphology, or how male and female reproductive traits covary among divergent populations. Here we address both topics using natural populations of the guppy, Poecilia reticulata, a livebearing fish that exhibits divergent patterns of male sexual behaviour among populations. Initially, we performed a series of mating trials on a single population to examine the relationship between the morphology of the male's copulatory organ (the gonopodium) and the success of forced matings. Using a combination of linear measurements and geometric morphometrics, we found that variation in the length and shape of the gonopodium predicted the success of forced matings in terms of the rate of genital contacts and insemination success, respectively. We then looked for geographical divergence in these traits, since the relative frequency of forced matings tends to be greater in high-predation populations. We found consistent patterns of variation in male genital size and shape in relation to the level of predation, and corresponding patterns of (co)variation in female genital morphology. Together, these data enable us to draw tentative conclusions about the underlying selective pressures causing correlated patterns of divergence in male and female genital traits, which point to a role for sexually antagonistic selection.     

Copulation behaviour of Glossina pallidipes (Diptera: Muscidae) outside and inside the female, with a discussion of genitalic evolution

If species-specific male genitalia are courtship devices under sexual selection by cryptic female choice, then species-specific aspects of the morphology and behaviour of male genitalia should often function to stimulate the female during copulation. The morphology and behaviour of the complex, species-specific male genitalia of the tsetse fly, Glossina pallidipes Austen, were determined from both direct observations and dissections of flash-frozen copulating pairs; we found that some male genitalic traits probably function to stimulate the female, while others function to restrain her. The male clamps the ventral surface of the female's abdomen tightly with his powerful cerci. Clamping does not always result in intromission. Clamping bends the female's body wall and her internal reproductive tract sharply, posteriorly and dorsally, and pinches them tightly. The male performed sustained, complex, stereotyped, rhythmic squeezing movements with his cerci that were not necessary to mechanically restrain the female and appeared instead to have a stimulatory function. Six different groups of modified setae on and near the male's genitalia rub directly against particular sites on the female during squeezing. The designs of these setae correlate with the force with which they press on the female and the probable sensitivity of the female surfaces that they contact. As expected under the hypothesis that these structures are under sexual selection by female choice, several traits suspected to have stimulatory functions have diverged in G. pallidipes and its close relative, G. longipalpis. Additional male non-genitalic behaviour during copulation, redescribed more precisely than in previous publications, is also likely to have a courtship function. The elaborate copulatory courtship behaviour and male genitalia may provide the stimuli that previous studies showed to induce female ovulation and resistance to remating.     

Male and female genitalia in some Libyan species of Acrididae (Orthoptera: Acridoidea)

A comparative study of male and female genitalia was carried out in thirty‐seven Libyan species representing twenty genera of the family Acrididae. An attempt has been made to describe and illustrate the different structures, namely, epiphallus, aedeagus, subgenital plate, supra‐anal plate and cerci of the male, and spermatheca, ovipositor, subgenital plate, supra‐anal plate and cerci of the female, in Acridids, with an aim to discover their significance in order to make the identification of genera and species, together with other generic characters, more perfect and convenient. Distinct family characters are shield or bridge‐shaped condition of epiphallus; presence or absence of dorso‐lateral appendices, oval sclerites and lophi on epiphallus; divided, undivided or flexured condition of aedeagus; presence or absence of gonopore process on aedeagus; condition of apical and pre‐apical diverticula of spermatheca; presence or absence of glandular pouches of Cornstock and Kellog on female subgenital plate; and rudimentary or well developed condition of egg‐guide. Stable characters for separating the subfamilies are taken to be presence or absence of ancorae on epiphallus, long or short condition of aedeagal sclerites; elongate, slender or short and broad condition of ovipositor valves: presence or absence of Jannone's organs and setae on posterior margin of female subgenital plate; and shape of diverticula of spermatheca. Useful generic characters are shape of male subgenital plate, supra‐anal plate and cerci, broad or narrow condition of bridge, presence or absence of branch of bridge connecting lophi with bridge of epiphallus; mono‐, bi‐ or tri‐lobate condition of lophi of epiphallus, length and upcurved or downcurved condition of apical valve of aedeagus, shape of posterior margin of female subgenital plate, presence of setae on the whole posterior margin or confined to lateral margins only; and toothed, tuberculate or smooth condition of ovipositor valves, length of the lateral apodeme in relation to the dorsal valves. Specific characters are shape of egg‐guide of female subgenital plate, shape of ovipositor valves and apical tips, shape of male supra‐anal plate and cerci, size of anterior and posterior lobes of lophi of epiphallus, size and shape of ancorae, shape of apical valves of aedeagus; and size of apical and pre‐apical diverticula and presence of protuberance on pre‐apical diverticulum.     

New investigation technique for thoracic sclerites of mosquitoes (Diptera, Culicidae) for the correct identification of genera and species

A new technique for studying the external morphology of thoracic sclerites of mosquitoes (Diptera: Culicidae) was developed. According to this method, the shape of sclerites and the position of setae and scales can be examined using SEM or light microscopy even in cases when setae or scales have been lost. The method can be recommended for the damaged material which is often obtained during sampling. The bases of setae usually exceed 100 μm in diameter and thus differ significantly from the bases of scales which are about 30 μm in diameter. Analysis of the structure of sclerites and the arrangement of setae and scales on them in different specimens of one species may reveal intraspecific variation of this character complex. Comparison of species from different genera of the family Culicidae, as well as comparison of closely related species, will probably help to estimate the diagnostic value of thoracic sclerites as morphological characters.