Andrology of shortnose guitarfish Zapteryx brevirostris (Müller & Henle, 1841) (Chondrichthyes, Trygonorrhinidae).

The andrological study of a species involves the macro- and microscopic analyses of the internal reproductive organs and the evaluation of seminal parameters and ultrastructural characteristics of the spermatozoa. As in other vertebrates, the male reproductive tract in chondrichthyans consists of testes and reproductive ducts (efferent duct, epididymis, Leydig's gland, ductus deferens and seminal vesicle). In this study the authors used three adult specimens of Zapteryx brevirostris from wild capture kept at the Ubatuba Aquarium, Brazil. Semen was collected by abdominal massage over the location of the seminal vesicle, preceded by ultrasonographic evaluation. The semen collected was diluted 1:200 and subject to quantitative and morphological analyses. Ultrastructural analysis was performed using transmission and scanning electron microscopy. Correlation was observed between successful collection and ultrasonographic image of an engorged seminal vesicle, as well as testicles with easily delimitable margins and higher echogenicity. It was possible to identify free spermatozoa with helical filiform appearance, as well as spermatozeugmata. The average sperm concentration resulted in 5 million packets per millilitre and 140 million spermatozoa per millilitre. The sperm nucleus is described as follows: cone shaped, parachromatin sheath less dense than the chromatin of the nucleus, smooth depression of the nuclear fossa, abaxial axoneme 9 + 2 and accessory axonemal columns in positions 3 and 8 and oval shaped, with flattened inner surface in cross-section. These results broaden the knowledge of the andrology of this species, contributing to ex situ breeding programmes.

Size, shape, and direction matters: Matching secondary genital structures in male and female mites using multiple microscopy techniques and 3D modeling.

Studies of female genital structures have generally lagged behind comparable studies of male genitalia, in part because of an assumption of a lower level of variability, but also because internal genitalia are much more difficult to study. Using multiple microscopy techniques, including video stereomicroscopy, fluorescence microscopy, low-temperature scanning electron microscopy (LT-SEM), and confocal laser scanning microscopy (CLSM) we examined whether the complex sperm transfer structures in males of Megalolaelaps colossus (Acari: Mesostigmata) are matched by similarly complex internal structures in the female. While both LT-SEM and CLSM are well suited for obtaining high-quality surface images, CLSM also proved to be a valuable technique for observing internal anatomical structures. The long and coiled sperm transfer organ on the chelicera of the males (spermatodactyl) largely matches an equally complex, but internal, spiral structure in the females in shape, size, and direction. This result strongly suggests some form of genital coevolution. A hypothesis of sexual conflict appears to provide the best fit for all available data (morphology and life history).

Penetration mechanics of elongated female and male genitalia of earwigs.

We unveiled the penile penetration mechanics of two earwig species, Echinosoma horridum, whose intromittent organ, termed virga, is extraordinarily long, and E. denticulatum, whose virga is conversely short. We characterised configuration, geometry, material and bending stiffness for both virga and spermatheca. The short virga of E. denticulatum has a material gradient with the stiffer base, whereas the long virga of E. horridum and the spermathecae of both species are homogeneously sclerotised. The long virga of E. horridum has a lower bending stiffness than the spermatheca. The virga of E. denticulatum is overall less flexible than the spermatheca. We compared our results to a previous study on the penetration mechanics of elongated beetle genitalia. Based on the comparison, we hypothesised that the lower stiffness of the male intromittent organ comparing to the corresponding female structure is a universal prerequisite for the penetration mechanics of the elongated intromittent organ in insects.

Motility of efferent duct cilia aids passage of sperm cells through the male reproductive system.

Motile cilia line the efferent ducts of the mammalian male reproductive tract. Several recent mouse studies have demonstrated that a reduced generation of multiple motile cilia in efferent ducts is associated with obstructive oligozoospermia and fertility issues. However, the sole impact of efferent duct cilia dysmotility on male infertility has not been studied so far either in mice or human. Using video microscopy, histological- and ultrastructural analyses, we examined male reproductive tracts of mice deficient for the axonemal motor protein DNAH5: this defect exclusively disrupts the outer dynein arm (ODA) composition of motile cilia but not the ODA composition and motility of sperm flagella. These mice have immotile efferent duct cilia that lack ODAs, which are essential for ciliary beat generation. Furthermore, they show accumulation of sperm in the efferent duct. Notably, the ultrastructure and motility of sperm from these males are unaffected. Likewise, human individuals with loss-of-function DNAH5 mutations present with reduced sperm count in the ejaculate (oligozoospermia) and dilatations of the epididymal head but normal sperm motility, similar to DNAH5 deficient mice. The findings of this translational study demonstrate, in both mice and men, that efferent duct ciliary motility is important for male reproductive fitness and uncovers a novel pathomechanism distinct from primary defects of sperm motility (asthenozoospermia). If future work can identify environmental factors or defects in genes other than DNAH5 that cause efferent duct cilia dysmotility, this will help unravel other causes of oligozoospermia and may influence future practices in genetic and fertility counseling as well as ART.

Coevolution between female seminal receptacle and sperm morphology in the semiaquatic measurer bug Hydrometra stagnorum L. (Heteroptera, Hydrometridae).

The coevolution between sperm length and size of the female sperm-storage organs is described for the first time within Heteroptera. The long sperm of the measurer bug Hydrometra stagnorum is characterized by the unusually long acrosome with its anterior region helically arranged, and by a very short nucleus. The sperm flagellum has a 9 + 9+2 conventional axoneme and crystallized mitochondrial derivatives. The female spermatheca consists of an extraordinarily long spermathecal duct ending with an apical spermathecal bulb into which flows also the secretions of a relatively short spermathecal gland. Both spermathecal duct and gland have a thin epithelium lined by a cuticle, beneath which a complex of secretory and duct forming cells are present. The secretions of these two structures flow into the apical spermathecal bulb. A thick layer of muscle fibers surrounds the epithelium. These results confirm the opinion that the dimensions of the female reproductive sperm-storage organs are able to drive the sperm morphology.

Ultrastructure and histochemistry of the male reproductive system of the genus Callinectes Stimpson, 1860 (Brachyura: Portunidae).

We described the ultrastructure and histochemistry of the reproductive system of five Callinectes species, and evaluate the seasonal variation in weight of the reproductive system and hepatopancreas by comparing annual changes of somatic indices. The somatic indices changed little throughout the year. In Callinectes, spermatogenesis occurs inside the lobular testes and, within each lobule, the cells are at the same developmental stage. Spermatogenesis and spermiogenesis follow the same development pattern in all Callinectes studied. Mature spermatozoa are released into the seminiferous ducts through the collecting ducts. Cells of the vas deferens are secretory as evidenced by rough endoplasmic reticulum, Golgi complex, and secretory vesicles that produce the seminal fluid. The anterior vas deferens shows two portions: proximal and distal. In proximal portion (AVDp), spermatozoa are clustered and embedded in an electron-dense, basophilic glycoproteinaceous secretion Type I. In the distal portion (AVDd), the spermatophore wall is formed by incorporation of a less electron-dense glycoproteinaceous secretion Type II. The secretion Type I change to an acid polysaccharide-rich matrix that separates the spermatophores from each other. The median vas deferens (MVD) stores the spermatophores and produces the granular glycoproteinaceous seminal fluid. The posterior vas deferens (PVD) has few spermatophores. Its epithelium has many mitochondria and the PVD seminal fluid changes into a liquid and homogeneous glycoprotein. Many outpocketings in the PVD and MVD help to increase the fluid production. Overall, the reproductive pattern of Callinectes is similar to other species that produce sperm plugs. The secretions of AVD, MVD, and PVD are responsible for the polymerization that forms the solid, waxy plug in the seminal receptacle. The traits identified here are common to all Portunidae species studied so far.

The male reproductive accessory glands of the blister beetle Meloe proscarabaeus Linnaeus, 1758 (Coleoptera: Meloidae): Anatomy and ultrastructure of the cantharidin-storing organs.

Blister beetles owe their name to their ability to release cantharidin, a blistering terpene, the highest concentration of which is retained in male accessory glands. The anatomy and ultrastructure of the three pairs of male reproductive accessory glands and the glandular region of the two vasa deferentia of Meloe proscarabaeus were investigated using light, electron and ion beam microscopy. All of the mesodermal glands here analysed share a common structural organization with an outer muscular layer and an inner glandular epithelium facing a broad lumen in which the secretory products are released. Developed rough endoplasmic reticulum, Golgi systems, abundant mitochondria, numerous secretory vesicles and a microvillated apical membrane are commonly found in the cells of different glandular epithelia, suggesting that all accessory gland pairs as well as the vasa deferentia are involved in an active synthesis. Nevertheless, each pair of glands appears specialized in the production of a specific set of substances, as suggested by the peculiarities in cellular ultrastructure and by the different aspect of the secretions stored in their glandular lumen. The above cited features of male accessory glands of M. proscarabaeus are compared with those of other beetles and some hints on their potential role in producing and/or concentrating cantharidin are provided.

Vasa deferentia and associated structures of the male Panorpodes kuandianensis (Mecoptera: Panorpodidae).

The male reproductive system may provide significant evidence for the taxonomic and phylogenetic analyses of insects. However, current knowledge of the male reproductive system in Mecoptera is mainly concentrated on the external genitalia, and is rarely involved in the internal reproductive system. Here, we investigated the morphology and the fine structure of the vasa deferentia and associated structures of the male reproductive system of Panorpodes kuandianensis Zhong et al., 2011 (Panorpodidae) using light, scanning, and transmission electron microscopy. The male reproductive system of P. kuandianensis consists of a pair of symmetrical testes with three tubular testicular follicles, two epididymides, two distinctly partitioned vasa deferentia, a pair of mesadenia, one ejaculatory sac, and the external genitalia. A pair of expanded seminal vesicles are modified from the median part of the vasa deferentia, and evolve into secretory organs. The seminal vesicles have elongated cylindrical epithelial cells, which contain abundant secretory materials in the cytoplasm and form a small central lumen, likely serving a secretory function rather than provisionally storing sperm as in most other insects. Alternatively, the sperm are stored temporarily in the epididymis, the greatly coiled portion of the vasa deferentia. The morphology of the male reproductive system supports the close relationships of Panorpidae and Panorpodidae.

Genital morphology and the mechanics of copulation in the millipede genus Pseudopolydesmus (Diplopoda: Polydesmida: Polydesmidae).

Mate choice, copulation, genital morphology, and sperm storage are not very well understood in millipedes. The use of three-dimensional x-ray computed tomography (µCT) provides new morphological data regarding millipede reproductive systems in both the female and male, including chitinous sclerites and membranes, muscles, glands, oviducts, and sperm conduits. Here we present a complete integrated account of the morphology and function of the female genital organs in the family Polydesmidae (Diplopoda: Polydesmida) using µCT, UV fluorescence imaging, and scanning electron microscopy. These data allow us to consider competing hypotheses regarding millipede vulva formation. We additionally present the morphology of copulatory interface in Pseudopolydesmus Attems, 1898 using images of a mating pair in copula and by simulating the interface of the organs using 3D models from µCT, allowing us to tentatively identify a lock-and-key-like mechanism. Finally, we use µCT to reveal the topology of the seminal canal in the gonopod of male Pseudopolydesmus, a topic that has remained unresolved for nearly 80 years.

Accessory glands of male reproductive system in Pseudochorthippus parallelus parallelus (Zetterstedt, 1821) (Orthoptera: Acrididae): A light and electron microscopic study.

The accessory glands of male reproductive system in insects play a significant role in the reproduction process by protecting sperm in spermatheca, preventing female to accept other males after mating and stimulating oviposition. The number, structure, and arrangement of the tubules of accessory glands can change from species to species. In this study, the accessory glands belonging the male reproductive system in Pseudochorthippus parallelus parallelus (Zetterstedt, 1821) (Orthoptera, Acrididae) were examined with stereomicroscope, light microscope, scanning (SEM), and transmission (TEM) electron microscopes at Gazi University, Faculty of Science in 2017-2019. P. parallelus parallelus is a widespread species that is located at the extending areas from Italy to the Northern Europe and also in Turkey. The accessory glands of P. parallelus parallelus' male reproductive system are composed of about 10 tubules. The tubules can be classified into two groups according to the thickness of their muscle tissues. Both groups have single layered epithelial cells with mitochondria, well-developed endoplasmic reticulum, spherical nucleus with electron dense chromatin, secretory vesicles and multivesicular bodies in their cytoplasm. In addition, apocrine type secretion is seen in epithelial cells.

Scanning Electron Microscopy of Sand Flies of the Chagasi Series, Psychodopygus (Diptera: Psychodidae) Genus, Focusing on the Genitalia.

Males of cryptic or closely related species present great morphological variation in their genitalia, whereas females, such as those of the Chagasi Series of the Psychodopygus Mangabeira, 1941 genus, are more similar. Therefore, our aim was to study the fine structure of the male genitalia of five species of the Chagasi Series to better understand the variation in their morphology and its influence on the copulatory process. The sand fly species were captured in the following Brazilian states: Psychodopygus chagasi (Costa Lima, 1941) (Rondônia), Psychodopygus complexus (Mangabeira, 1941) (Tocantins), Psychodopygus squamiventris maripaensis (Floch & Abonnenc, 1946) (Amapá), Psychodopygus squamiventris squamiventris (Lutz & Neiva, 1912) (Amazonas), and Psychodopygus wellcomei Fraiha, Shaw & Lainson, 1971 (Pará and Ceará). Insects were stored in ethanol 70% (then dehydrated) and dry after they were sputtered with gold. The samples were observed under a scanning electron microscope. Microtrichiae, two types of trichoid sensilla, coeloconic and chaetic sensillae, were observed on the antenna of all species, with no difference between them. Only on the anepimeron of P. squamiventris squamiventris a modified 'racket'-like scale was observed. As for the male genitalia, the setae and structures of each species were fully described, such as the small setae on the paramere apex of the P. squamiventris subspecies, and the grooves present in this region and on the paramere lobe of P. complexus and P. wellcomei, which are impossible to observe with optic microscopy. New information is thus provided on the male genitalia, which can contribute to future bionomic studies of these species.

New Anatomical Evidence from the Male Asian Citrus Psyllid (Hemiptera: Liviidae) Invokes Controversy Over the Accepted Function of Some Male Reproductive Organs in Psylloidea.

Males of many Psylloidea are known to possess a characteristic structure at the functional hub of their reproductive apparatus, between afferent and efferent passage of seminal fluid. The structure is a squat, cylindrical endoskeleton consisting of two sections. Classical authors named them as 'sperm pump' and 'ejaculatory duct', based on superficial resemblance to a spring-loaded, thimble-shaped cylinder, encircled by smooth, vertical columns interpreted to be muscles which, when contracted, compress the cylinder and affect seminal fluid discharge. The discovery of numerous spherules of unknown composition and function in and around the columns of the Asian citrus psyllid male genitalia invoked rigorous scrutiny of the classical literature for evidence to support its claims, and determined that the grounds for vetting the structure as a sperm pump were fully teleological. This paper raises several objections to modern acceptance of this classical interpretation, presenting them as problematic, thought-provoking, and sometimes controversial anatomical features. The two sections are herein called 'drum' and 'spout'. As an endoskeleton, the sections are an invagination of the exoskeleton and therefore cannot receive seminal fluid into their hollow. A phallus is identified inside an aedeagal tube, indicating that it is the ejaculatory duct-the tube, drum, and spout are considered its housing. A sheath envelopes the drum and is directly continuous with the spout hypodermis, another problematical feature raising the question of whether it is detached from adherence to the drum cuticles. Also, there are four afferent tubes but only two openings in the drum to receive their seminal fluids.

The sperm pump and genital coupling of Panorpodes kuandianensis (Mecoptera: Panorpodidae).

Males of Panorpodidae possess a special sperm pump, through which they directly transfer seminal fluid to the female spermatheca. However, the sperm pump has not been studied in Panorpodes to date. Here, the structure of the sperm pump and the internal coupling of genitalia were investigated in the short-faced scorpionfly Panorpodes kuandianensis Zhong, Zhang, and Hua, 2011 using light and scanning electron microscopy. The sperm pump mainly consists of a piston, a pumping chamber, the anterior region of the aedeagal complex, the posterior region of the ejaculatory sac, and associated muscles. The piston as a propulsion apparatus is controlled by levator and depressor muscles. Its posterior region connects dorsally to the aedeagus via a joint. The pumping chamber is located between the piston and the aedeagus. The dorsal and ventral parameres were attached by retractor muscles. During copulation, the male phallotreme connects to the female copulatory pore to transfer sperm. Male gonostyli and parameres grasp the female to restrict the genitalia movement and impede her medigynium from retreating. The sperm ejaculatory mechanism of Panorpodes and the evolution of sperm transfer mode in insects are briefly discussed based on the structure of the sperm pump and the internal coupling of genitalia.

Genital morphology and copulatory behavior in triatomine bugs (Reduviidae: Triatominae).

Triatomines (Heteroptera: Reduviidae) include around 139 species, widely known as vectors of Chagas disease. Our aim is to review the existing knowledge of the genital morphology and sexual behavior and provide some functional analysis of these traits in triatomines. A complex set of traits comprise genitalia and these are highly variable among species. The components of the phallus and seminal products (secreted by action of testes and two accessory glands) interact to allow successful sperm transfer to the female spermathecae (usually a pair of blind tubes that emerge from the common oviduct). Seminal products may inhibit female physiology and extend mating duration. Mating behavior in triatomines is best characterized as scramble competition. We suggest that males may evaluate female condition prior to copulation, given that female fitness is largely affected by food (blood) source. Although rearing several triatomine species may be difficult and discourage from undertaking studies on this group, any further investigation on sexual behavior and mating interactions may provide data for applicative studies including Chagas disease vectors control.

Traumatic insemination is not the case in three Orius species (Heteroptera: Anthocoridae).

Traumatic insemination (TI) is an extraordinary style of mating behavior wherein the female integument is pierced by the male extragenital structure to transfer the spermatozoa into the female's body through wounding. Flower bugs of the genus Orius belong to the family Anthocoridae (Heteroptera), which is referred to as the "TI family". Males possess sharp shaped extragenitalia, and females receive the extragenitalia using the copulatory tubes, which are specialized extragenital structures in Orius species. Since TI is not well studied in insects possessing the copulatory tube, we examined the genital structures and copulatory processes of three species, Orius strigicollis, O. sauteri, and O. minutus. Scanning electron microscopic observations revealed the positions of male extragenital structures during copulation. A needle-like flagellum was deeply inserted into the female intersegment between the abdominal VII and VIII segments, while the curved part of a sickle-like cone forced the intersegment to expand. No scars were detected around the copulation region after copulation. The copulatory tube adhered to the interior of segment VII, and the interior integument around the copulatory tube remained intact after copulation. On the basis of these results, TI does not occur in these Orius species. A pair of seminal conceptacles, which exists in typical TI insects, was found at the base of the oviducts in O. strigicollis. The distal end of the copulatory tube connected to a closed bag with a double-membrane, termed the sperm pouch. The sperm pouch was filled with filamentous structures after copulation and structures with equivalent forms were observed in adult male testis. These structures, considered to be spermatozoa, persisted in the pouch for at least two weeks after copulation, suggesting that the pouch is a long-term spermatozoa storage organ.

Copulatory mechanism and functional morphology of genitalia and anal horn of the scorpionfly Cerapanorpa dubia (Mecoptera: Panorpidae).

The scorpionfly genus Cerapanorpa is characterized by the male possessing a single finger-like anal horn on the posterior portion of tergite VI. However, the functional morphology of this anal horn and the genitalia have not been studied to date. Herein, we investigated the functional morphology of the genitalia and the nongenital structures of the scorpionfly Cerapanorpa dubia by observing the mating process and dissecting the freeze-fixated pairs in copula to reveal the copulatory mechanism. The male C. dubia provides a solid salivary mass to the female as a nuptial gift prior to copulation. When the female starts to feed on the gift, the male uses his notal organ and complex genital structures to control the closest wing and genitalia of the female to establish a V-shaped mating position. In the maintenance phase of copulation, the male uses his anal horn in cooperation with the basally-constricted abdominal segment VII to clamp female abdominal segment VIII. The male hypovalves grasp female cerci, and move up and down rhythmically. The paired parameres clasp both sides of female tergite IX. The basal processes on male gonostyli grip the pleural membranes of the female genital chamber. In the sperm transfer phase, the male aedeagus directly couples with the female medigynium to transmit sperm by connecting his phallotreme to the female's copulatory pore. The evolution of the male complex grasping structures in Panorpidae is also briefly discussed.

The functional anatomy of external genitalia in the black tiger prawn (Penaeus monodon) studied by micro-computed tomography and scanning electron microscopy.

Although artificial insemination has been used for decades in Penaeus monodon aquaculture, the interaction of male and female external genitalia during spermatophore transfer has not been fully documented. As a result, studying the functional anatomy of this process may help to better refine the insemination technique. The sexual act in penaeoid prawns is virtually impossible to observe directly; as a result, this study aimed to describe the functional anatomy and interaction of external genitalia, such as the petasma, appendices masculinae and genital papillae of the male, with that of the thelycum and genital lobes of the female, using a combination of micro-computed tomography and scanning electron microscopy. We hypothesise that the spermatophores are ejaculated into the ventromedial groove of the petasma and squeezed by abdominal flexure into the spermathecae of female's thelycum. During this process, the 'spike-like' setae observed on the petasma and appendices masculinae are speculated to control the transferring direction of the spermatophores. The approach of three dimensional remodelling and animation reported in this study may prove useful in the examination of further hypotheses related to the functional anatomy of external genitalia and/or appendages for other crustacean species.

Morphological and histological structure of the male reproductive system of the water strider Gerris lacustris (Linnaeus 1758) (Gerridae, Heteroptera).

This work presents the male reproductive system morphology and histology of the water strider Gerris lacustris (Linnaeus 1758) (Gerridae, Heteroptera) using light microscopy and scanning electron microscopy. The male reproductive system of G. lacustris comprise of a pair of testes, two vasa deferentia, two seminal vesicles, an ejaculatory duct. There is no bulbus ejaculatorius and the long vas deferantia uniting to form a simple ductus ejaculatorius which is connected to the aedeagus. The testes are white colored and this cylindiric-shaped structure lies along genital abdominal segment. The testicular follicles have three different development zones (growth zone, maturation zone, differentiation zone). Each testis has two follicles, which are not lined by a common peritoneal sheath and involving many cysts arranged in a progressive order of maturation from the distal to the proximal region; spermiogenesis occurs in mature males, finishing with the organization of sperm bundles. The testes are connected to the seminal vesicles, specialized sperm storage places, by the vas deferentia.

Ebselen can Protect Male Reproductive Organs and Male Fertility from Manganese Toxicity: Structural and Bioanalytical Approach in a Rat Model.

Ebselen (EBS) is a versatile compound that can protect the cellular components from oxidative and free radical-mediated damage. In the present study, we investigated the protective effect of EBS against manganese (Mn) toxicity on male reproductive organs. Thirty-two male rats were assigned into four groups, namely, negative control, EBS (15 mg/kg body weight (bw), as a single protective IP injection), MnCl2 (50 mg/kg bw, orally for 30 consecutive days), and EBS + MnCl2 (as mentioned before). The results showed that EBS ameliorated the alterations caused by MnCl2 in the testicular, epididymal, and seminal vesicle tissues. MnCl2 increased the sperm abnormalities, decreased gonadosomatic index (GSI), sperm motility, and sperm count. Further, it reduced the serum levels of testosterone (T) and luteinizing hormone (LH). The elevated levels of malondialdehyde (MDA), nitric oxide (NO), and 8-OH-2'-deoxyguanosine (8-OHdG) and decreased the levels of superoxide dismutase (SOD), glutathione (GSH), and catalase (CAT) upon exposure to MnCl2 indicated a disturbance in the activities of the testicular antioxidant enzymes and indices. Histologically, MnCl2 decreased the diameter of seminiferous tubules (ST), the height of germinal epithelium, number of spermatogonia/ST, spermatocytes/ST, spermatids/ST, and Leydig cells/intertubular area (IA). Chemoprotection with EBS successfully mitigated most of the above-mentioned parameters concluding that EBS could be used as a useful prophylactic therapy whenever Mn toxicity is involved.

The morphology of the reproductive system in the crab Percnon gibbesi (Decapoda: Brachyura: Grapsoidea) reveals a new combination of characters in Thoracotremata.

Recent studies revealed a high diversity of reproductive structures in heterotreme brachyurans, while those of Thoracotremata seem rather uniform. Yet, there still is a huge lack of data in this group as only few species have been studied with respect to their reproductive system. The phylogenetic position of Percnidae is ambiguous. Recent molecular studies place it within polyphyletic grapsoids. We herein study the reproductive morphology of Percnon gibbesi using histology, scanning electron microscopy, micro-computed tomography and 3D-reconstructions to test whether this species shows the characteristic thoracotreme pattern. Our results reveal that the male copulatory system conforms to other thoracotremes. It is composed of two pairs of pleopods (gonopods) and likewise paired penes. The first gonopod is relatively long. It possesses a bent terminal process with a distal opening of the ejaculatory canal, a character also present in other grapsoids. The second gonopod is short and terminates in an apical girdle. The female reproductive system reveals a combination of characters, so far unknown for thoracotremes. The paired oviducts do not lead into the seminal receptacles, but run into separate cuticular ducts joined with the vaginae. Accessory sperm storage organs, the bursae, are also connected to the vaginae. Bursae have previously only been described in heterotreme crabs. The data presented in this study reveals a higher diversity of thoracotreme reproductive systems than anticipated.