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.     

Morphology and function of the reproductive system of representatives of the genus Uca

The morphology of the reproductive organs of three species of fiddler crabs, Uca ecuadoriensis, Uca c.f. forcipata, and Uca tangeri were investigated to subsequently produce a model of their mode of operation. Vulva, vagina, and spermatheca in females, and the first and second gonopods in males were examined by applying histological techniques and electron microscopy. In all three species, vulva and vagina conform to the concave type, and the spermatheca complies with the ventral type. The tissue of the oviduct orifice is enlarged and bulges into the lumen of the spermatheca. Differences between the three species are apparent in the organization of the spermatheca, especially in the distribution and structure of glandular epithelium: In U. ecuadoriensis and U. c.f. forcipata the largest proportion of the spermathecal wall is lined with cuticle and only a small area consists of glandular epithelium, while in U. tangeri almost all of the lining is glandular. Furthermore, the glandular epithelia of the species differ in their histology and ultrastructure: In U. ecuadoriensis it is tubular and multilayered, while in U. c.f. forcipata it is mono‐layered. U. tangeri finally has both forms of this tissue. In the males, the terminal segments of the first gonopod exhibit a tight fit to female organs and narrow, tightly sealed sperm channels. These features suggest a tendency towards minimizing loss of fluids, which can be interpreted as an adaptation to mating on land. The tight fit of male gonopod and female opening seem to be protection from interbreeding, which points toward a strong sexual selection. In the terrestrial environment, these originally aquatic organisms experience serious competition for resources; therefore there is pressure on successful reproduction. According to the current results a model of the process of fertilization and egg‐laying involving the investigated organs was generated. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Global diversity of crabs (Crustacea: Decapoda: Brachyura) in freshwater

An assessment of the global freshwater crab diversity is presented. A total of 1,476 species in 14 families are currently known from all zoogeographical regions (except Antarctica), including 1,306 species in eight exclusively freshwater families (Pseudothelphusidae, Trichodactylidae, Potamonautidae, Deckeniidae, Platythelphusidae, Potamidae, Gecarcinucidae and Parathelphusidae). Estimates of true freshwater crab diversity including likely numbers of undescribed taxa suggest that the field remains largely in a “discovery” phase. Main ideas on the origins, diversification, and phylogeny of true freshwater crabs are briefly discussed. The economic importance of freshwater crabs is also highlighted. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Relative growth and onset of morphological sexual maturity of the freshwater crab Sylviocarcinus pictus in a river of the Caatinga of northeastern Brazil

The aim of this study was to investigate the relative growth and onset of morphological sexual maturity of the freshwater crab Sylviocarcinus pictus. Specimens were collected every month from October 2013 to September 2014, at night, on a stretch of the river Guaribas, Piauí, Brazil. Crabs were sexed and their carapace width (CW, independent variable), carapace length (CL), cheliped propodus length (PL) and height (PH), gonopod length (GL), and abdomen width (AW) (dependent variables) were measured. These measurements were related to characterise relative growth and possible sex differences. On average males were larger than females (p = 0.0001). Size at the onset of morphological sexual maturity was defined by relating CL vs. PL for males (30.82 mm) and CL vs. AW for females (28.63 mm). These are considered secondary sexual characters and reflect initial size at morphological sexual maturity with greater precision. The differential cheliped growth of males may be related to courting and disputes with other males, while the allometric growth of the abdomen of females indicates increased probability of reproductive success.     

DNA taxonomy of the potamid freshwater crabs from Northern Africa (Decapoda,Potamidae)

The Mediterranean area, from southern Balkans to western Maghreb, is inhabited by the Potamon subgenus Euthelphusa, with three currently recognised species. The Maghrebian species P. (E.) algeriense is isolated from other Potamon species by the Mediterranean Sea and the Sahara Desert, and nearly no molecular data are currently available for this taxon. Present study investigated the mtDNA and nuDNA diversity in Potamon algeriense s.l. with the aims of exploring its molecular diversity pattern throughout its known distribution range, and testing the possible presence of cryptic taxa currently lumped under this binomen. The phylogenetic and DNA taxonomy analyses showed the presence of two well-supported clades of species rank within P. algeriense s.l, with an eastern clade including the populations from Tunisia and Numidia, and a western, highly structured clade including the populations from central Algeria and Morocco. In agreement with a typical Maghrebian biogeographic pattern, the distribution of these two species shows a clear east–west divide, with a disjunction zone located in Kabylia, and a strong link between molecular diversity and segregation within different hydrographical basins is evident. The Maghreb thus proved to host an unexpectedly high genetic diversity of, and to constitute a biodiversity hot-spot for, the Potamon subgenus Euthelphusa. In the light of the existence of two well-characterised species currently lumped under P. algeriense s.l., and of their noteworthy molecular structuring, the status of Maghrebian Potamon populations should be re-assessed for both the species present in the area, which are to be considered as independent management units.     

A comparative study of zoeal morphology in the genus Liocarcinus (Crustacea: Brachyura: Portunidae)

The zoeal morphology of Liocarcinus arcuatus, L. corrugatus and L. depurator is described from laboratory reared material and the larvae of L. holsatus, L. puber, L. pusillus and L. marmoreus are reexamined from specimens deposited in the BM (NH). No single meristic character was found that would adequately separate the seven described species of Liocarcinus. Characters that may facilitate separation of later zoeal stages only are listed.     

Morphology of the female reproductive system of three dorippid crabs (Crustacea: Decapoda: Brachyura: Dorippidae) and the role of accessory cuticle structures associated with seminal receptacles

The reproductive systems of crabs reveal characters of considerable importance for the understanding of brachyuran phylogeny and evolution. The Dorippoidea show several plesiomorphic characters within Eubrachyura and similarities to podotreme crabs. Hence, they are often considered as an early diverging lineage, sometimes even as the sister group to all remaining eubrachyurans. Due to their role as prime candidates for putative plesiomorphic characters of the reproductive system of the Eubrachyura, we compared the morphology of the vaginae, seminal receptacles, and ovaries of three dorippid species using histological methods, micro‐computed tomography, and 3‐D reconstructions. Despite the putative phylogenetic position of dorippids, the female reproductive system shows features that are regarded as derived characters in eubrachyurans, including a concave vagina and a ventral‐type seminal receptacle. In contrast to other eubrachyurans, the oviduct does not enter the seminal receptacle directly but through specific cuticular valves. The female reproductive systems of Dorippe sinica and Dorippe quadridens are remarkable in further aspects. The seminal receptacles of both species are completely cuticle‐lined and have accessory sperm storage structures, the bursae. Our findings on the morphology of the female reproductive system of dorippids with its unique combination of basal, derived, and new characters challenges the prevailing hypothesis on the evolution of sperm storage organs in Eubrachyura.     

Ultrastructure and relationships of spermatozoa of the freshwater crabs Potamon fluviatile and Potamon ibericum (Crustacea, Brachyura, Potamidae)

The spermatozoa of Potamon fluviatile and P. ibericum are virtually indistinguishable and do not support separate subgeneric rank ( Potamon and Pontipotamon , respectively). Synapomorphic with the spermatozoa of the South African freshwater crab Potamonautes perlatus Sidneyi are the elongation of the two centrioles and disposition of the centrioles almost parallel to each other, unknown elsewhere in the Brachyura, and reduction of the thickened ring (homoplasic with grapsids and gecarcinids). Other, probably synapomorphic, similarities of Potamon and Potamonautes include the wide inner acrosome zone, absence of a definite acrosome ray zone (homoplasic in other families) and the cleistospermial spermatophores. Further similarities, of questionable polarity, are the simple, not multilaminar, nuclear membrane and the tendency of the nuclear arms to wrap around the nucleus. Differences of Potamon from Potamonautes , which possibly support their present generic separation and give weak support to their former separate familial placement in the Potamidae and Potamonautidae respectively, are perforation of the operculum and the weak, rather than strong, development of a periopercular rim. Absence in Potamon and Potamonautes of an accessory opercular ring and a xanthid ring separates them from xanthids. No close affinities with other heterotreme families are seen but their assignment to the Helerotremata is not in doubt. Their spermatozoa lack two of the distinctive features of thoracotreme spermatozoa (apical button on the operculum and concentric lamellation of the outer acrosome zone). No clear correlates of spermatozoon structure with a freshwater existence are recognizable but reduction of the thickened ring possibly relates to peculiarities of the acrosome reaction. However, the production of spermatophores with single spermatozoa (cleistospermia) is possibly a device to prevent polyspermy and wastage of the small number of lecithotrophic eggs produced in potamids.     

Characteristics of the motor and genital copulatory responses of the male hamster

The temporal and dynamic characteristics (duration, frequency, rhythmicity, and vigor) of pelvic thrusting displayed by 17 male golden hamsters during their copulatory behavioral responses: mounts, intromissions, ejaculations, and long intromissions, as well as their temporal correlation with the genital contacts established by intravaginal penile insertion, were studied by an accelerometric and polygraphic technique. Pelvic thrusting in all copulatory behavioral responses appeared as series or trains of rhythmical, synchronic, vigorous movements. The various pelvic thrusting trains lasted around 1 s on average, but those of mounts were significantly longer than the trains displayed in the other behavioral responses. The frequencies of pelvic thrusting were similar in all responses (around 15 thrusts per s) excepting mounts which had lower values. In intromission, ejaculation, and long intromission responses, when penile insertion occurred, pelvic thrusting either was interrupted or showed changes in its characteristics: penile insertion was related to a period without thrusting in intromissions, to a series of intravaginal thrusting of higher frequency (16.4 thrusts per s) and lower vigor in ejaculations, and to a prolonged period of 6 to 25 s of slow intravaginal pelvic thrusting (1–2 thrusts per s) in long intromissions. Penile insertion lasted longer in ejaculations than in intromissions and it was significantly shorter in both of these responses than in long intromissions. These results provide information about some dynamic aspects of sexual behavior in hamsters, as well as a temporal correlation between the motor and genital components of this behavior.     

Functional morphology of the reproductive system of Galathea intermedia (Decapoda: Anomura)

Spermatophore formation in Galathea intermedia begins in the proximal part of the vas deferens. The contents subsequently form a spermatophoric ribbon, the so-called "secondary spermatophore," in its distal part. A strongly muscular ductus ejaculatorius is present in the coxa of the fifth pereiopod which builds up pressure for the extrusion of the spermatophoric ribbon. After extrusion, the ribbon is caught by the first gonopod, while the second gonopod dissolves the matrix of the ribbon. During copulation the spermatophores are randomly placed on the sternum of the female, near the genital opening, by the fifth pereiopods of the male. Subsequent ovulation of the female via the genital opening, an active process accomplished through muscular activity, results in fertilization of the eggs by the exploding spermatophores. External intersexes are characterized by both male and female external sexual characters, but in all individuals only male gonads are present. No trace of a female reproductive system could be detected. Thus, these external intersexes are exclusively functional males.     

Action of a rhizocephalan on the genital activity of host male crabs: characterization of a parasitic secretion inhibiting spermatogenesis

Protein fractions obtained by filtration of crude extracts of roots from the sacculinid parasite Loxothylacus panopei were tested by repeated injections on healthy male crabs Rhithropanopeus harrisii, the usual host of the rhizocephalan. Testes, and rogenic glands and different parts of the host CNS were observed; the behaviour of the animals was also noted. For the first time, proteinaceous substance(s) of about 25000–30000 daltons were characterized. They induce inhibition of spermatogenesis, and rogenic gland cytolysis and depletion of the sinus gland as in naturally infested crabs. The mode of action of sacculinid on spermatogenesis in the host is discussed.     

Ultrastructure of spermatozoa and spermatophores of old world freshwater crabs (Brachyura: Potamoidea: Gecarcinucidae,Potamidae, and Potamonautidae)

We investigated the ultrastructure of spermatozoa and spermatophores of 19 palaeotropical freshwater crab species [12 species of the Gecarcinucidae, 6 of the Potamidae (Potamiscinae), and 1 species of the Potamonautidae (Deckeniinae: Hydrothelphusini)]. The investigated Potamiscinae have densely packed coenospermic spermatophores with the exception of Thaiphusa sirikit and Johora singaporensis that exhibit cleistospermia. In contrast, in the Gecarcinucidae the spermatozoa are loosely embedded in a mucous matrix. The gecarcinucid and potamiscine sperm differ, furthermore, in acrosomal structure and size. The acrosome in the Gecarcinucidae is much smaller and spherical, while the larger acrosome in the Potamiscinae has the tendency to be depressed. In the Potamiscinae, an additional middle acrosomal zone evolved between the acrosome ray zone and the outer acrosomal zone. Within the Gecarcinucidae, a differentiation into two groups (Gecarcinucinae and Parathelphusinae) is not supported by the present spermatological data. The sperm morphology of Hydrothelphusa aff. madagascariensis (Potamonautidae: Deckeniinae) differs from Potamonautes sidneyi (Potamonautidae: Potamonautinae) in acrosomal size and shape, and in the absence of a periopercular rim. A closer relationship of Deckeniinae and Gecarcinucidae cannot be confirmed by spermatology. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

中国肩花蝽属雌性交配管的形态研究

半翅目花蝽科（Anthocoridae） 昆虫的种类鉴定通常是以雄性外生殖器特征为主, 雌虫的种类鉴定一直是研究中的难点。本文研究了中国肩花蝽属 Tetraphleps 已知5个种雌虫的交配管,包括黑色肩花蝽T. aterrimus (Sahlberg)、斑翅肩花蝽T. galchanoides Ghauri、毛肩花蝽T. pilosulus Bu et Zheng、直长肩花蝽T. parallelus Bu et Zheng和玉龙肩花蝽T. yulongensis Bu et Zheng。发现交配管的构造在种内变异小、特征稳定,而在种间差别明显,可作为该属雌虫种类鉴定的可靠依据。     

The copulatory plug delays ejaculation by rival males and affects sperm competition outcome in house mice

Females of many species mate with multiple males (polyandry), resulting in male–male competition extending to post‐copulation (sperm competition). Males adapt to such post‐copulatory sexual selection by altering features of their ejaculate that increase its competitiveness and/or by decreasing the risk of sperm competition through female manipulation or interference with rival male behaviour. At ejaculation, males of many species deposit copulatory plugs, which are commonly interpreted as a male adaptation to post‐copulatory competition and are thought to reduce or delay female remating. Here, we used a vertebrate model species, the house mouse, to study the consequences of copulatory plugs for post‐copulatory competition. We experimentally manipulated plugs after a female's first mating and investigated the consequences for rival male behaviour and paternity outcome. We found that even intact copulatory plugs were ineffective at preventing female remating, but that plugs influenced the rival male copulatory behaviour. Rivals facing intact copulatory plugs performed more but shorter copulations and ejaculated later than when the plug had been fully or partially removed. This suggests that the copulatory plug represents a considerable physical barrier to rival males. The paternity share of first males increased with a longer delay between the first and second males' ejaculations, indicative of fitness consequences of copulatory plugs. However, when males provided little copulatory stimulation, the incidence of pregnancy failure increased, representing a potential benefit of intense and repeated copulation besides plug removal. We discuss the potential mechanisms of how plugs influence sperm competition outcome and consequences for male copulatory behaviour.     

On the phylogenetic significance of sagittocysts and copulatory organs in acoel turbellarians

Viewed by SEM and TEM, sagittocysts of Convoluta bifoveolata Mamkaev, 1971, and needles of C. sagittifera Ivanov, 1952, have the same structure. Both are capsule-form extrusomes containing a protrusible needle. Only seven similar species of convolutimorph acoels symbiotic with green algae and C. sagittifera, without algae, possess extrusomes of this peculiar and complicated type. The sagittocyst is a clear synapomorphy of all these species. A sacciform ciliated antrum lacking a seminal vesicle is also characteristic of these species and also of three Japanese species of green (algae-symbiotic) convolutimorph acoels lacking sagittocysts. We suggest schemes of the possible evolution of male and female copulatory organs to provide a basis for better using such organs as phylogenetic characters. We regard the formation of a ciliated sacciform antrum as an independent evolutionary trend. This conclusion forms the basis for establishing the separate family Sagittiferidae. Species of this family seem to have originated in the West Pacific.     

Corpus al latum regulation of copulatory behaviour in the male grasshopper, Melanoplus sanguinipes

Abstract The relationship between corpus allatum (CA) regulation of copulatory behaviour and the CA's influence on the development of the accessory reproductive glands was studied in the male grasshopper, Melanoplus sanguinipes (Fabr.). Allatectomy slowed the development of copulatory behaviour; ≤80% of control males 7 days and older mated with females, but allatectomized males did not show comparable levels of copulation until day 19. In a 3 h observation period, the time taken for 50% of males to initiate copulation was c. 60 min for allatectomized males and only c. 10 min for controls. At the age when 50% of males copulated, the accessory glands of allatectomized males were less developed than those of controls; the accessory gland weight was 65%, soluble protein was 72% and reserves of long hyaline protein I were 40% those of controls. Treatment with Juvenile Hormone III on day 2 stimulated accumulation of long hyaline protein I in 9-day-old allatectomized males to the level found in untreated 19-day-old allactec-tomized males, but did not elicit heightened copulation. No positive feedback from the accessory glands appeared necessary to elicit copulation since vasectomy, ablation of the accessory glands or isolation of the terminal abdominal ganglion by ventral nerve cord transection did not inhibit copulatory behaviour in 7-day-old males. Because allatectomy inhibited copulatory behaviour in 7-day-old males in which the accessory glands were removed but not in 19-day-old males without accessory glands, the increased tendency of older allatectomized males to mate with females was not due to release of inhibition caused by continued growth of the accessory reproductive glands.     

Comparative ultrastructure of the spermatozoa of the Majoidea (Crustacea,Decapoda, Brachyura) with new data on six species in five genera

Comparative ultrastructure of majoid spermatozoa belonging to 23 species, in 19 genera and five families, is considered, with new data on Schizophrys aspera; S. rufescens (Majidae, Majinae); Camposcia retusa (Inachidae); Pyromaia tuberculata (Inachoididae); and Huenia heraldica and Menaethius monoceros (Epialtidae, Epialtinae). The oregoniid Chionoecetes opilio, and inachids Cyrtomaia furici, Platymaia rebierei, Macropodia longirostris and Inachus phalangium, possibly with Camposcia retusa, but not Podochela riisei, appear to form a group. Within the inachids, Macropodia and Inachus are especially close. A domed central acrosome zone, seen in most inachid sperm, in majines (both Schizophrys species), in pisines (Oxypleurodon orbiculatus and O. stuckiae) and epialtines (Huenia heraldica and Menaethius monoceros), appears to be an autapomorphy of these majoids. A peripheral acrosome zone is seen in the inachid Grypacheus hyalinus, two inachoidids (P. tuberculata and Stenorhynchus seticornis) and the majid Maja squinado. Pyromaia tuberculata differs from other inachoidids in having a slightly dome‐shaped operculum. The mithracine Macrocoeloma trispinosum (Majidae) sperm more closely resembles Inachoididae, than Inachidae. Spermatologically, the family Majidae and the subfamily Majinae are not homogeneous. Spermatozoal ultrastructure does not support a majoid–hymenosomatid relationship and is equivocal with regard to the placement of Cryptochiridae in either the Thoracotremata or Heterotremata, the prominent operculum strongly differentiates cryptochirids from Majoidea.     

Long-term persistence of male copulatory behavior in castrated and photo-inhibited Siberian hamsters

Gonadal steroids are essential for the long-term maintenance of the full repertoire of sexual behavior in male rodents. Typically, all individuals of several species cease to display the ejaculatory reflex within a few weeks of castration. The present study documents the persistence of the ejaculatory reflex 19 weeks after orchidectomy in 40% of male Siberian hamsters maintained in long or short day lengths; testosterone was undetectable in the circulation of these animals. Intact hamsters transferred from a long to a short photoperiod underwent gonadal regression: 50% of these animals continued to display mating behavior culminating in ejaculation throughout 25 weeks of testing. The remaining animals failed to ejaculate after approximately 11 weeks of short day treatment but resumed mating coincident with spontaneous gonadal recrudescence. Activation of sex behavior in the latter cohort appears to depend on gonadal steroids and is in contrast to the copulatory behavior of the substantial proportion of the study population that sustains the full sexual repertoire in the long-term absence of gonadal steroids. Sex behavior of the latter animals may be dependent on nongonadal steroids or mediation by steroid-independent mechanisms.