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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.

Alpacas (Vicugna pacos) are reported to be the rare mammal in which the penis enters the uterus in mating. To date, however, only circumstantial evidence supports this assertion. Using female alpacas culled for meat, we determined that the alpaca penis penetrates to the very tips of the uterine horns, abrading the tract and breaking fine blood vessels. All female alpacas sacrificed one hour or 24 hours after mating showed conspicuous bleeding in the epithelium of some region of their reproductive tract, including the hymen, cervix and the tips of each uterine horn, but typically not in the vagina. Unmated females showed no evidence of conspicuous bleeding. Histological examination of mated females revealed widespread abrasion of the cervical and endometrial epithelium, injuries absent in unmated females. Within one hour of mating, sperm were already present in the oviduct. The male alpaca's cartilaginous penis tip with a hardened urethral process is likely responsible for the copulatory abrasion. The entire female reproductive tract interacts with the penis, functioning like a vagina. Alpacas are induced ovulators, and wounding may hasten delivery of the seminal ovulation-inducing factor beta-NGF into the female's blood stream. There is no evidence of sexual conflict in copulation in alpaca, and thus wounding may also be one of a variety of mechanisms devised by mammals to induce a beneficial, short-term inflammatory response that stimulates blastocyst implantation, the uterine remodeling associated with placental development, and thus the success of early pregnancy., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Brennan et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

What is the vagina? This seemingly simple question has a rather complex answer, depending on whether we use a functional or a developmental definition. The terminal portion of the female reproductive tract that opens to the environment initially served as a conduit for eggs to be laid, and in species with external fertilization the distal oviduct may be specialized for oviposition but there is no vagina. In animals with internal fertilization, this terminal section of the oviduct interacts with the sperm and the intromittent organ leading to functional specialization of this region that we often call a vagina in insects and some vertebrates. Here we address the evolution, morphology and diverse functions of the vagina and some of the unknown questions that remain to be addressed in the study of this remarkable structure., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

Female genitalia are conspicuously overlooked in comparison to their male counterparts, limiting our understanding of sexual reproduction across vertebrate lineages. This study is the first complete description of the clitoris (hemiclitores) in female snakes. We describe morphological variation in size and shape ( n = 9 species, 4 families) that is potentially comparable to the male intromittent organs in squamate reptiles (hemipenes). Dissection, diffusible iodine contrast-enhanced micro-CT and histology revealed that, unlike lizard hemiclitores, the snake hemiclitores are non-eversible structures. The two individual hemiclitores are separated medially by connective tissue, forming a triangular structure that extends posteriorly. Histology of the hemiclitores in Australian death adders ( Acanthophis antarcticus ) showed erectile tissue and strands/bundles of nerves, but no spines (as is found in male hemipenes). These histological features suggest the snake hemiclitores have functional significance in mating and definitively show that the hemiclitores are not underdeveloped hemipenes or scent glands, which have been erroneously indicated in other studies. Our discovery supports that hemiclitores have been retained across squamates and provides preliminary evidence of differences in this structure among snake species, which can be used to further understand systematics, reproductive evolution and ecology across squamate reptiles.

While most birds do not have external genitalia, in a small number of species the males have penises, which can be either intromittent or non-intromittent. Brennan provides an overview of bird genitalia, including speculation about the evolutionary forces involved., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Despite their evolutionary and biomedical importance, studies of the morphology and function of female genitalia have continued to lag behind those of male genitalia. While studying female genitalia can be difficult because of their soft, deformable and internal nature, recent advances in imaging, geometric analyses of shape and mechanical testing have been made, allowing for a much greater understanding of the incredible diversity of form and function of female genitalia. Here we summarize some of these methods, as well as discuss some big questions in the field that are beginning to be examined now, and will continue to benefit from further work, especially a comparative approach. Topics of further research include examination of the morphology of female genitalia in situ, in-depth anatomical work in many more species, studies of the interplay between natural and sexual selection in influencing features of vaginal morphology, how these diverse functions influence the mechanical properties of tissues, and studies of clitoris morphology and function across amniotes. Many other research topics related to female genitalia remain largely unexplored, and we hope that the papers in this issue will continue to inspire further research on female genitalia., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

The lower reproductive tract of female mammals has several competing functions including mating, tract health maintenance, and parturition. Diverse vaginal anatomy suggests interactions between natural and sexual selection, yet despite its importance, female copulatory morphology remains under-studied. We undertook a comparative study across the species-rich mammalian order Chiroptera (bats) with a focus on the suborder Yangochiroptera (Vespertilioniformes) to examine how female vaginal features may have coevolved with male penis morphology to minimize mechanical damage to their tissues during copulation. The penis morphology is diverse, presenting great potential for post-copulatory sexual selection and coevolution with the female morphology, but vaginas have not been carefully examined. Here we test the hypotheses that vaginal thickness and collagen density have coevolved with features of the male penis including the presence of spines and a baculum. We present histological data from females of 24 species from 7 families of bats, and corresponding data on male penis anatomy. We also examine the role of phylogenetic history in the morphological patterns we observe. We found evidence that female vaginal thickness has coevolved with the presence of penile spines, but not with baculum presence or width. Collagen density did not appear to covary with male penile features. Our findings highlight the importance of considering interactions between the sexes in influencing functional reproductive structures and examine how these structures have been under selection in bats., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Genital evolution can be driven by diverse selective pressures. Across taxa we see evidence of covariation between males and females, as well as divergent genital morphologies between closely related species. Quantitative analyses of morphological changes in coevolving male and female genitalia have not yet been shown in vertebrates. This study uses 2D and 3D geometric morphometrics to quantitatively compare the complex shapes of vaginal pouches and hemipenes across three species of watersnakes (the sister taxa Nerodia fasciata, N. sipedon, and a close relative N. rhombifer) to address the relationship between genital morphology and divergence time in a system where sexual conflict may have driven sexually antagonistic coevolution of genital traits. Our pairwise comparisons of shape differences across species show that the sister species have male and female genitalia that are significantly different from each other, but more similar to each other than to N. rhombifer. We also determine that the main axes of shape variation are the same for males and females, with changes that relate to deeper bilobation of the vaginal pouch and hemipenes. In males, the protrusion of the region of spines at the base of the hemipene trades off with the degree of bilobation, suggesting amelioration of sexual conflict, perhaps driven by changes in the relative size of the entrance of the vaginal pouch that could have made spines less effective., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Zoos and natural history museums are both collections-based institutions with important missions in biodiversity research and education. Animals in zoos are a repository and living record of the world's biodiversity, whereas natural history museums are a permanent historical record of snapshots of biodiversity in time. Surprisingly, despite significant overlap in institutional missions, formal partnerships between these institution types are infrequent. Life history information, pedigrees, and medical records maintained at zoos should be seen as complementary to historical records of morphology, genetics, and distribution kept at museums. Through examining both institution types, we synthesize the benefits and challenges of cross-institutional exchanges and propose actions to increase the dialog between zoos and museums. With a growing recognition of the importance of collections to the advancement of scientific research and discovery, a transformational impact could be made with long-term investments in connecting the institutions that are caretakers of living and preserved animals., (© The Author(s) 2022. Published by Oxford University Press on behalf of the American Institute of Biological Sciences.)

In species that copulate during non-conceptive periods, such as humans and bonobos, sexual intercourse is known to be pleasurable for females. Dolphins also copulate throughout the year, largely to establish and maintain social bonds 1 . In dolphins, the clitoris is positioned in the anterior aspect of the vaginal entrance 2 , where physical contact and stimulation during copulation is likely. Clitoral stimulation seems to be important during female-female sexual interactions in common bottlenose dolphins (Tursiops truncatus), which rub each other's clitorises using snouts, flippers, or flukes 3 . Determining a sexual pleasure response in animals not amenable to neurobehavioral examination is difficult, but investigation of the clitoris may elucidate evidence of functionality. In this study, we assessed macro- and micromorphological features of the clitoris in common bottlenose dolphins to examine functional features, including erectile bodies with lacunae, extensible collagen and/or elastin fibers, and the presence and location of sensory nerves. Our observations suggest the clitoris of dolphins has well-developed erectile spaces, is highly sensitive to tactile stimulation, and is likely functional. VIDEO ABSTRACT., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Copulatory behavior and genital morphology interact to deliver sperm more effectively during mating, but the nature of this interaction has not been explored in depth in most vertebrates. Alpacas have unusually long copulations lasting 15-20 min, and a unique copulatory behavior, where the penis intromits all the way past the cervix, into the uterine horns. Here we describe the morphology of male and female genitalia and report unique morphological characteristics that may be associated with this unusual insemination mode. Vaginal shape is highly variable, and seemingly not associated with age or parity. The cranial vagina varies between bulbous and cylindrical, while the caudal vagina is typically narrower and always cylindrical. The cervix consists of a series of two to three spirals or rings, and it is often found in a relaxed state that may prevent damage caused by the cartilaginous penis tip as it pushes through the cervix to reach the uterine horns. The uterus and uterine horns have a complex shape with multiple constrictions. The cartilaginous penis tip has a sharp urethral process that may help to push against these constrictions. The diameter of the vaginal lumen is much greater than the diameter of the penis suggesting that there is little direct interaction between them, and that female vaginal shape is not under strong copulatory selection. In effect, the entire female reproductive tract of the female is interacting with the penis during copulation., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Comparisons of 3D shapes have recently been applied to diverse anatomical structures using landmarking techniques. However, discerning evolutionary patterns can be challenging for structures lacking homologous landmarks. We used alpha shape analyses to quantify vaginal shape complexity in 40 marine mammal specimens including cetaceans, pinnipeds, and sirenians. We explored phylogenetic signal and the potential roles of natural and sexual selection on vaginal shape evolution. Complexity scores were consistent with qualitative observations. Cetaceans had a broad range of alpha complexities, while pinnipeds were comparatively simple and sirenians were complex. Intraspecific variation was found. Three-dimensional surface heat maps revealed that shape complexity was driven by invaginations and protrusions of the vaginal wall. Phylogenetic signal was weak and metrics of natural selection (relative neonate size) and sexual selection (relative testes size, sexual size dimorphism, and penis morphology) did not explain vaginal complexity patterns. Additional metrics, such as penile shape complexity, may yield interesting insights into marine mammal genital coevolution. We advocate for the use of alpha shapes to discern patterns of evolution that would otherwise not be possible in 3D anatomical structures lacking homologous landmarks., Competing Interests: None declared., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

The phallic glans of the American alligator (Alligator mississippiensis) is the distal termination of the semen-conducting sulcus spermaticus and during copulation has the closest, most intimate mechanical interactions with the female urodeum, the middle cloacal chamber that contains the opening to the vaginal passages and oviducts. However, the details of this interface leading to insemination and gamete uptake are unclear. Here, we: (1) histologically characterize the underlying tissue types and morphologically quantify the shape changes associated with glans inflation into the copulatory conformation, (2) digitally reconstruct from MRI the 3D shape of functional tissue compartments, and (3) diffusible iodine-based contrast-enhanced computed tomography image the copulatory fit between male phallus and female cloaca. We discuss these results in relation to tissue type material properties, the transfer on intromittent forces, establishing potential copulatory lock, inflated glans volume scaling with body mass/length, the mechanics of semen targeting and insemination, and potential female cryptic choice impacting multiple clutch paternity. In part, this study further clarifies the phallic morphological variation observed among crocodylians and begins to investigate the role(s) these divergent male forms play during copulation interacting with female cloacal forms to increase reproductive success., (© The Author(s) 2020. Published by Oxford University Press on behalf of Society for the Study of Reproduction. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Asymmetric genitalia and lateralized mating behaviors occur in several taxa, yet whether asymmetric morphology in one sex correlates or coevolves with lateralized mating behavior in the other sex remains largely unexplored. While lateralized mating behaviors are taxonomically widespread, among mammals they are only known in the harbor porpoise (Phocoena phocoena). Males attempt copulation by approaching a female exclusively on her left side. To understand if this unusual lateralized behavior may have coevolved with genital morphology, we quantified the shape of female and male harbor porpoise reproductive tracts using 2D geometric morphometrics and 3D models of the vaginal lumen and inflated distal penis. We found that the vaginas varied individually in shape and that the vaginas demonstrated both significant directional and fluctuating asymmetry. This asymmetry resulted from complex 3D spirals and vaginal folds with deep recesses, which may curtail the depth or direction of penile penetration and/or semen movement. The asymmetric shapes of the vaginal lumen and penis tip were both left-canted with similar angular bends that mirrored one another and correspond with the left lateral mating approach. We suggest that the reproductive anatomy of both sexes and their lateral mating behavior coevolved.

Whales, dolphins, and porpoises have unusual vaginal folds of unknown function(s) that are hypothesized to play an important role in sexual selection. The potential function of vaginal folds was assessed by testing the mechanical properties of common bottlenose dolphin (Tursiops truncatus) reproductive tract tissues in 6 different regions and across age classes in post-mortem specimens. We assessed the regional (local) and overall effective elastic modulus of tissues using indentation and tensile tests, respectively. We explore the non-linear mechanical response of biological tissues, which are not often quantified. Indentation tests demonstrated that sexual maturity state, tissue region, force history, and force magnitude values significantly affected the measured effective elastic modulus. Tissue was stiffest in the vaginal fold region and overall stiffer in sexually immature compared to mature animals, likely reflecting biomechanical adaptations associated with copulation and parturition. Tensile tests showed that only tissue region significantly affected the effective modulus. Our data support the hypothesis that vaginal folds function as mechanical barriers to the penis and may provide females with mechanisms to reduce copulatory forces on other reproductive tissue. STATEMENT OF SIGNIFICANCE: Cetaceans have unusual folds of vaginal wall tissue that appear to evolve under sexual selection mechanisms and present physical barriers to the penis during copulation. We explore the biomaterial properties of vaginal fold tissue, how it varies from other reproductive tract tissues, and ontogenetic patterns. We demonstrate that vaginal folds can withstand higher mechanical forces and respond in a manner conducive to dissipating copulatory forces to other reproductive tissues. This study yields exciting insights on how female genital tissue may function during copulation, and is the first to do so in any vertebrate species. Additionally, we provide an example for testing biological tissues, non-linear properties, and materials with uneven surface structure and uneven thickness., (Copyright © 2019 Acta Materialia Inc. Published by Elsevier Ltd. All rights reserved.)

Male genital diversification is likely the result of sexual selection. Female genital diversification may also result from sexual selection, although it is less well studied and understood. Female genitalia are complex among whales, dolphins, and porpoises, especially compared to other vertebrates. The evolutionary factors affecting the diversity of vaginal complexity could include ontogeny, allometry, phylogeny, sexual selection, and natural selection. We quantified shape variation in female genitalia using 2D geometric morphometric analysis, and validated the application of this method to study soft tissues. We explored patterns of variation in the shape of the cervix and vagina of 24 cetacean species (n = 61 specimens), and found that genital shape varies primarily in the relative vaginal length and overall aspect ratio of the reproductive tract. Extensive genital shape variation was partly explained by ontogenetic changes and evolutionary allometry among sexually mature cetaceans, whereas phylogenetic signal, relative testis size, and neonate size were not significantly associated with genital shape. Female genital shape is diverse and evolves rapidly even among closely related species, consistent with predictions of sexual selection models and with findings in invertebrate and vertebrate taxa. Future research exploring genital shape variation in 3D will offer new insights into evolutionary mechanisms because internal vaginal structures are variable and can form complex spirals., (© 2017 The Author(s). Evolution © 2017 The Society for the Study of Evolution.)

Genitalia are morphologically variable across many taxa and in physical contact during intromission, but little is known about how variation in form correlates with function during copulation. Marine mammals offer important insights into the evolutionary forces that act on genital morphology because they have diverse genitalia and are adapted to aquatic living and mating. Cetaceans have a fibroelastic penis and muscular vaginal folds, while pinnipeds have a baculum and lack vaginal folds. We examined copulatory fit in naturally deceased marine mammals to identify anatomical landmarks in contact during copulation and the potential depth of penile penetration into the vagina. Excised penises were artificially inflated to erection with pressurized saline and compared with silicone vaginal endocasts and within excised vaginas in simulated copulation using high-resolution, diffusible iodine-based, contrast-enhanced computed tomography. We found evidence suggestive of both congruent and antagonistic genital coevolution, depending on the species. We suggest that sexual selection influences morphological shape. This study improves our understanding of how mechanical interactions during copulation influence the shape of genitalia and affect fertility, and has broad applications to other taxa and species conservation., (© 2017 The Author(s).)