Morphology of the male reproductive system of freshwater prawn Macrobrachium carcinus (Decapoda,Caridea): Functional and comparative aspects

Testes and vasa deferentia are parts of the male reproductive system of decapod crustaceans. Both organs show morphological differences among decapod species in terms of anatomical and histological patterns reflecting the diversity of this group. Describing these features may assist in systematics, phylogenetics, and studies of reproductive behavior, especially for species of commercial interest, such as Macrobrachium carcinus, a native American species that, unusually for this genus, has no precopulation courting behavior. This study aims to describe the reproductive morphology and spermatogenesis of the male freshwater prawn M. carcinus. The male reproductive system of this species consisted of lobed testes connected to the vasa deferentia. The testis of M. carcinus was divided into several lobules. Each lobule was formed by a cluster of germ cells surrounded by connective tissue and nurse cells. This microscopic anatomy and histology of the testicular histoarchitecture has been described for many species of Decapoda and may represent a derived design of the testes. Unlike that in other decapod species, spermatogenesis proceeds in short transitory phases that produce spermatozoa at high concentrations and frequencies, corroborating the uncommon male reproductive behavior of this species. In the spermatic pathway, the lobules develop and fuse before releasing spermatozoa from the testes; however, this process has not been observed in decapods, yet. The neutral compounds secreted by the vas deferens are important for sperm nutrition as females secrete a substance for spermatophore adhesion during reproduction. This study presents different features and dynamics of the spermatogenic process in the male reproductive system of M. carcinus that have not yet been presented in the literature for decapods.     

Spermatophore transfer in the hermit crab Clibanarius vittatus (Crustacea,Anomura, Diogenidae)

Although mating has been described in several hermit crab species, the mechanics of spermatophore transfer have not previously been demonstrated. Evidence from pleopod and gonopore morphology, video observations, and inseminated females indicates that in Clibanarius vittatus the male applies a spermatophoric mass directly onto the female via the gonopores rather than with modified pleopods 1-2 (gonopods) and/or genital papillae as in many other decapods. The single second pleopod of males of C. vittatus has a simple endopod with no apparent modifications for sperm transfer. There are no genital papillae extending from the male gonopores. The globular spermatophores are aligned in rows surrounded by a seminal secretion in the male ducts (vasa deferentia that terminate in ejaculatory ducts opening to the exterior via the gonopores). During copulation, described from time-lapse video recordings, the ventral surface of the last thoracic segment of the male, bearing the gonopores, was apposed to the ventral cephalothorax of the female. A massive amount of seminal secretion containing spermatophore ribbons, termed here the spermatophoric mass and described for the first time in a hermit crab species, was observed covering the sternites and coxae of pereopods 1-5 of a recently copulated female. It is suggested that during copulation the male emits the contents of the ejaculatory ducts directly onto the female without the aid of gonopods or genital papillae. Although spermatophore transfer is simple in C. vittatus, the presence of modified anterior pleopods or elongate genital papillae (sexual tubes) in other paguroidean species suggests the possibility of a more complex insemination process in these other hermit crabs.     

Dimorphic ejaculates and sperm release strategies associated with alternative mating behaviors in the squid

Sperm competition is a powerful postcopulatory selective force influencing male adaptations associated with increasing fertilization success, and it is usually related to the evolution of different strategies of ejaculate expenditure between individuals. Ejaculates may also be influenced by additional selective pressures associated with sperm competition, such as timing between insemination and fertilization, female reproductive tract morphology, and fertilization environment. Also, males that adopt alternative mating tactics may face distinct sperm competition pressures, which may lead to the evolution of intraspecific diversity in ejaculates. In loliginid squids, males with alternative reproductive tactics (sneakers and consorts) differ not only in mating behavior, but also transfer spermatophores into two distinct sites within the female. Here, we compared structure and functioning of spermatophores between sneakers and consorts in the squid Doryteuthis plei applying microscopy techniques and in vitro experiments. Sneakers and consorts exhibit differences in spermatophore structure that lead to distinct spermatophoric reactions and spermatangium morphologies. Moreover, in sneakers, sperm release lasts longer and their sperm show an aggregative behavior not detected in consorts. Slow sperm release may be a strategy to guarantee longer sperm provision, given the wide interval between sneaker mating and egg release. For consorts, in turn, intense and quick sperm discharge may be advantageous, as timing between mating and egg‐laying is relatively short. Within the complex squid mating system, factors such as (i) different fertilization sites and (ii) interval between mating and egg release may also influence sperm competition, and ultimately shape the evolution of divergent ejaculates between dimorphic males.     

The functional morphology and role of the thelycum in insemination,and its relation to the mating system in the seagrass shrimp Ambidexter symmetricus (Decapoda: Processidae)

There is considerable variation in structures known to function in the transfer and storage of sperm in female decapod crustaceans. The thelycum is a secondary sexual character that forms from the posterior thoracic sterna of female shrimps (especially penaeoids and sergestoids). Females in the caridean shrimp family Processidae have a thelycum‐like structure which rarely occurs in other caridean females. We tested the hypothesis that the processid thelycum serves as a spermatheca for either short‐term attachment or long‐term sperm storage. When inseminated females of the processid Ambidexter symmetricus were isolated after mating, newly spawned and then incubated embryos hatched, but in the continued absence of a male, females were unable to fertilize a subsequent spawn. Our observations on A. symmetricus show that sperm were not retained after female spawning, and thus the thelycum is not used for long‐term sperm storage as in many penaeoids. In A. symmetricus, the thelycum may serve as an external median spermatheca (seminal receptacle) for temporary attachment and storage of a sperm mass during the 2–3 h interval between mating and spawning. Observations on mating behavior support the hypothesis of a pure‐search (promiscuous) mating system in A. symmetricus, with males showing little interest before, and copulating with females only after, the female parturial molt. Mating encounters were short (<2 min). This mating system is like that of other caridean shrimps with populations structured similarly to those in A. symmetricus: a relatively high density of mobile individuals and sexual dimorphism in body size (reproductive females larger than males) but not in cheliped weaponry (similar in males and females).     

The reproductive system of Osedax (Annelida,Siboglinidae): ovary structure,sperm ultrastructure,and fertilization mode

Osedax is a genus of siboglinid annelids in which the females live on dead vertebrate bones on the seafloor. These females have a posterior end that lies within the bone and contains the ovarian tissue, as well as the “roots” involved with bone degradation and nutrition. The males are microscopic and live as “harems” in the lumen of the gelatinous tube that surrounds the female trunk, well away from the ovary. Females are known to spawn fertilized primary oocytes, suggesting internal fertilization. However, little is known about sperm transfer, sperm storage, or the location of fertilization, and the morphology of the female reproductive system has not been described and compared with the reproductive systems of other siboglinids. A 3D‐reconstruction of the ovisac of Osedax showed ovarian tissue with multiple lobes and mature oocytes stored in a “uterus” before being released through the single oviduct. The oviduct emerges as a gonopore on the trunk and travels along the trunk to finally open to the seawater as a thin cylindrical tube among the crown of palps. Light and transmission electron microscopy of mature Osedax sperm revealed elongate heads consisting of a nucleus with helical grooves occupied by mitochondria. In contrast to other Siboglinidae, Osedax sperm are not packaged into spermatophores or spermatozeugmata, and Osedax females lack a discrete region for sperm storage. Transmission electron microscopy and fluorescence microscopy allowed detection of sperm associated with ovarian tissue of the female ovisac of four different Osedax species. This provides the first evidence for the site of internal fertilization in Osedax. A heart body was found in the circulatory system, as seen in other siboglinids and some other annelids. The possible presence of nephridia in the anterior ovisac region was also documented. These morphological features provide new insights for comparing the regionalization of Osedax females in relation to other siboglinids.     

虾蟹精子发育的营养及代谢研究概况

对雄性虾蟹的生殖营养及精子代谢作了综述。大量研究证实,在人工养殖条件下,雄性虾蟹的生殖力会出现明显下降,其原因与人工饲养条件下营养缺乏有关。精子储存在贮精囊和纳精囊中时,不断进行旺盛的代谢活动,其代谢类型主要是无氧代谢,而代谢的能量物质大部分种类为碳水化合物,这些物质主要由精子以外的物质提供。     

Chonopeltis australis (Crustacea) male reproductive system morphology; Sperm transfer and review of reproduction in Branchiura

The morphology of the male reproductive system as well as sperm transfer in Branchiura has been described for Dolops ranarum and Argulus japonicus. In this study, the reproductive system and accessory structures are described for male Chonopeltis australis using histology, light microscopy, and scanning electron microscopy. For the first time, we describe sperm transfer by means of a spermatophore in this genus. The internal and external morphology and mechanism of sperm transfer is compared with other Branchiura, where it has been described. The morphology of the reproductive system of C. australis is similar to that of D. ranarum while the accessory structures and the spermatophore produced are similar to that of A. japonicus. A revision of the definition of Branchiura with respect to reproduction is provided. J. Morphol. 276:209–218, 2015. © 2014 Wiley Periodicals, Inc.     

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.     

Sperm Quantity Explains Age-Related Variation in Fertilization Success in the Hide Beetle

Age-related variation in sperm quality and quantity may have a dramatic impact on female fecundity and fertilization success. Despite this, the relationships between these parameters are rarely investigated. Moreover, studies exploring age-related variation in male mating success generally fail to consider the entire lifespan of an individual male; instead they restrict analyses to a small number of defined age classes. As a consequence they are unable to assess the impact of early and late life history components on male reproductive success. In this study, we explore these questions using the hide beetle, Dermestes maculatus , a model species for investigating age-related male mating success. We first employed a longitudinal design to explore whether patterns of sperm transfer and subsequent reproductive success varied over the reproductive lives of a cohort of males. Secondly, we investigated age-related variation in sperm viability, a surrogate measure of sperm quality. Finally, we combined these data and assessed whether the observed patterns of sperm transfer were correlated with fertilization success. We found that the quantity of sperm varied with male age: the amount of sperm transferred to a female increased with male age until 9 wk and then started to decline. Similarly, female fecundity and fertilization success were related to male age: females mating with males when they were relatively young (1 wk) or relatively old (13 wk) suffered reductions in fecundity and fertilization. Our data suggest that fertilization success is driven at least in part by the quantity of sperm transferred during mating.     

Internal anatomy and ultrastructure of the male reproductive system of the Norway lobster Nephrops norvegicus (Decapoda: Astacidea)

The Norway lobster (Nephrops norvegicus) is economically important in Europe. However, apart from the female reproductive system, very little is known about its internal anatomy. This article focuses on studying the internal anatomy and ultrastructure of the male reproductive system. This system follows the general pattern found among decapod crustaceans, with several peculiarities. Testes are composed of lobular sperm ducts in which the spermatozoa are fully constituted. The spermatozoa present three lateral arms and a long acrosome, which gives a false appearance of flagellated spermatozoa. The two testes form a double H under the heart, and the vas deferens (VD) arise from each side at the posterior edge of the double H. The main characteristic of the VD is the presence of a sphincter in the enlarged area of the distal end of the middle VD. The MVD here shows an increase in musculature of the wall as compared to the VD, which regulates the passage of the sperm cord to the distal VD (DVD) and thence to the thelycum of the female. The wall of the spermatophore is formed in the distal part of the proximal VD, which surrounds the unique sperm cord present in the VD. Isolated spermatophores are not observed in the VD. The sperm cord is pinched off during copulation by the musculature of the DVD. Then, a portion of the sperm cord is transferred from each VD to form the isolated spermatophores. The wall of the spematophores and the spermatozoa that are observed inside the thelycum have the same morphology as those observed in the VD. J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

Masculinization of Female Crayfish,Procambarus Clarki (Girard)

Summary

Twenty-three immature female crayfish (Procambarus clarki (Girard); carapace length = 8.2–17.9 mm), implanted with androgenic gland implants obtained from P. clarki males, were observed for 326 days for evidence of masculinization. Number of molts completed ranged from three to eight. Of eleven surviving females, three were masculinized as evidenced by the partial transformation of their first pair of pleopods into male-type gonopods. No other evidence of masculinization, either internal or external was seen. Masculinization of the first pair of pleopods was unequal and first noted 290 days post-implantation. We conclude that androgenic gland implantation can induce masculinization of P. clarki females, albeit only with great difficulty. This is the first report of masculinization of female macruran decapods by androgenic gland implantation.     

Mating games squid play: reproductive behaviour and sexual skin displays in Caribbean reef squid Sepioteuthis sepioidea

Observation of the sexual interactions of Sepioteuthis sepioidea squid during the short reproductive stage of their lives showed a scramble competition system, with both male and female polygyny. Mature females were faithful to a specific location in the daytime, whereas males moved from group to group and formed short-term consortships with females. Males defended females from other males, particularly with an agonistic Zebra display. Male–female pairs exchanged Saddle-Stripe displays, after which males might display an on–off Flicker. There was considerable female choice. Only if a female responded to this display with a parallel Rocking action would she pair and would the male deposit spermatophores at the base of her arms, and only 50% of the time did females move the spermatophores internally to where sperm might be released and stored in the oviducal gland for later fertilization of eggs. This long-term set of interactions and solitary deposition of hidden egg strings contrasts with the attraction of both sexes to a common ‘egg mop’ laid by many females which was a site of competition in other loliginid squid. Since Sepioteuthis is a primitive genus within the family Loliginidae, it may represent a generalist reproductive strategy that evolved into a specialized localization one.     

Mate Choice by Non-Virgin Females Contributes to Reproductive Isolation between Populations of the Harlequin Beetle-Riding Pseudoscorpion

Determining whether reproductive isolation evolves through mate choice and/or gametic factors that prevent fertilization or through the post‐zygotic mechanisms of hybrid sterility or inviability is fundamental to understanding speciation. Investigation of the pre‐ and post‐zygotic components of reproductive isolation is facilitated in the pseudoscorpion, Cordylochernes scorpioides, by its indirect method of sperm transfer and viviparous embryonic development. Previous research on this species, in which mate discrimination was assessed in virgin females, suggested that female choice played only a minor role in reproductive isolation between populations from French Guiana and Panamá. Here, in a study of three allopatric populations of C. scorpioides from Panamá, we assessed mating‐stage isolation in both virgin and once‐mated females, and found that female discrimination depends critically on mating status. Virgin females were almost invariably receptive, showing no tendency to discriminate against males from allopatric populations. By contrast, non‐virgin females were significantly more likely to reject foreign males than males from their own population. Male sexual motivation could not account for differences in either female sexual receptivity or male success in sperm transfer. Allopatric and sympatric males did not differ in number of spermatophores deposited as either a female’s first or second mate. Nonetheless, allopatric males achieved significantly lower sperm transfer success not only with choosy, non‐virgin females but also with virgin females. Given the lack of behavioral discrimination by virgin females, female receptivity was not the only factor influencing differences in sperm transfer success. Multivariate analysis of spermatophore morphology suggests that the higher failure rate of matings between allopatric males and virgin females resulted from population differences in sperm packet architecture. Overall, our findings indicate that assessment of discrimination against allopatric males that is limited to virgin females may seriously underestimate the contribution of female mate choice to reproductive isolation between populations.     

Gonopore development and sex change in the Antarctic shrimp Chorismus antarcticus (Caridea: Hippolytidae)

Chorismus antarcticus is the only protandrous hermaphrodite decapod species found on the Antarctic continental shelves. The morphological structures of the male and female gonopores were described and used to determine sex and to fix the size range at which the transition from male to female takes place. The parallel occurrence of male and female gonopores was found in all specimens. The presence of open gonopore flaps at both the third and fifth pair of pereiopods in individuals between 9 and 12 mm of carapace length (CL) is discussed as the morphological indicator of sex change. Few females occurred from 9 mm CL onwards, whereas all shrimps larger than 13 mm CL only had open female gonopores. This size corresponds exactly with the CL at which onset of oocyte development and female gonad production starts. Secondary sexual characters, the gonopore structures, seem to allow an accurate prediction of the size range at which hermaphrodite caridean decapods change sex. Received: 23 February 1996/Accepted: 15 July 1996     

Female control of sperm ejection and retention in the cornsilk fly Euxesta eluta (Diptera: Ulidiidae)

Promiscuous mating systems provide the opportunity for females to bias fertilization toward particular males. However, distinguishing between male sperm competition and active female sperm choice is difficult for species with internal fertilization. Nevertheless, species that store and use sperm of different males in different storing structures and species where females are able to expel all or part of the ejaculates after copulation may be able to bias fertilization. We report a series of experiments aimed at providing evidence of female sperm choice in Euxesta eluta (Hendel), a species of ulidiid fly that expels and consumes ejaculates after copulation. We found no evidence of greater reproductive success for females mated singly, multiply with the same male, or mated multiply with different males. Female E. eluta possesses two spherical spermathecae and a bursa copulatrix for sperm storage, with a ventral receptacle. There was no significant difference in storing more sperm in spermathecae 24 h after copulation than immediately after copulation. Females mated with protein-fed males had greater reproductive success than similar females mated to protein-deprived males. Protein-fed females prevented to consume the ejaculate, retained more sperm when mated to protein-fed males than when mated to protein-deprived males. Our results suggest that female E. eluta can exert control of sperm retention of higher quality males through ejaculate ejection.     

Postmating sexual selection: the effects of male body size and recovery period on paternity and egg production rate in a water strider

The role of male body size in postmating sexual selection wasexplored in a semiaquatic insect, the water strider Gerris lateralis.To separate effects of male size per se from those due to numericsperm competition, male recovery period (shown here to be proportionalto ejaculate size) was manipulated independently of body sizein a factorial experiment where virgin females were mated firstwith sterile males and then with focal males. Both relativemale fertilization success and female reproductive rate were measured.The number of sperm transferred increased with male recoveryperiod, an effect that was mediated by longer copulation duration,but there were no effects of body size on ejaculate size. Neithermale size nor recovery period had any significant direct effectson male fertilization success. However, copulation durationinfluenced relative fertilization success, suggesting that malesable to transfer more sperm also achieved higher fertilizationsuccess. Females exercised cryptic female choice by modulatingtheir reproductive rate in a manner favoring large males andmales that were successful in terms of achieving high relativefertilization success. Thus, successful males gained a twofoldadvantage in postmating sexual selection. This study has important implicationsfor previous estimates of sexual selection in this group of insectsbecause pre- and postmating sexual selection will be antagonisticdue to limitations in male sperm production: males mating frequently(high mating success) will on average transfer fewer sperm ineach mating and will hence tend to fertilize fewer eggs permating (low fertilization success).     

Morphology and histology of male and female reproductive systems in the inseminating species Scoloplax distolothrix (Ostariophysi: Siluriformes: Scoloplacidae)

The morphology and histology of male and female reproductive systems were examined in Scoloplax distolothrix. Internal insemination was documented in this species by the presence of sperm within the ovaries. Mature males and females have elongated genital papillae, exhibiting a tubular shape in males and a plain heart‐shape with two median protuberances in females. The testes are two elongated structures that converge ventrally, under the intestine, towards the genital papilla. They are joined at the caudal end, forming an ovoid single chamber for sperm storage. Secretory regions were not observed. In the lumen of the testicular tubules, spermatozoa can be tightly packed along their lengths, but do not constitute a spermatozeugmata. The lumen of the sperm storage chamber and spermatic duct are filled with free spermatozoa without the accompanying secretions. The ovaries are bird‐wing shaped, saccular structures that converge ventrally under the intestine, towards the genital papilla. They are joined at the caudal end, forming a tubular chamber possibly destined for oocyte storage. An oviduct with an irregular outline connects the chamber to the tubular region of the genital papilla. No distinct sperm storage structure was found in the ovaries. The unique male and female genital papillae suggest that these structures are associated with the reproductive mode in scoloplacids, representing evidence for insemination. The occurrence of free spermatozoa, without the accompanying secretions and not arranged in a spermatozeugmata can be associated with the presence of a tubular male genital papilla for sperm transfer to the female genital tract. This reinforces the idea that sperm packets are not necessary for all inseminating species. The male reproductive system in scoloplacids is very different from that in auchenipterids, a second catfish family with insemination, which indicates that the occurrence of insemination is not connected to the internal morphology of reproductive organs. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Male and female effects on sperm precedence in the giant sperm species <Emphasis Type="Italic">Drosophila bifurca</Emphasis>

Sperm competition is expected to be a driving force in sexual selection. In internally fertilized organisms, it occurs when ejaculates from more than one male are present simultaneously within the female’s reproductive tract. It has been suggested that greater sperm size may improve the competitive ability of sperm, but studies provide contradictory results depending on the species. More recently, the role of females in the evolution of sperm morphology has been pointed out. We investigate here the male and female effects that influence sperm precedence in the giant sperm species, Drosophila bifurca Patterson & Wheeler. Females were mated with two successive males, and the paternity outcomes for both males were analyzed after determining sperm transfer and storage. We found very high values of last male sperm precedence, suggesting a strong interaction between rival sperm. However, the data also indicate high frequencies of removal of the sperm of the first male from the female reproductive tract prior to any interaction with the second male. This implies that successful paternity depends mainly on successful sperm storage. Knowing what happens to the sperm within females appears to be a prerequisite for disentangling post-copulatory sexual interactions between males and females.     

Multiple mating,sperm transfer and oviposition pattern in the giant sperm species,Drosophila bifurca

Many diverse traits are involved in gamete systems, and several models have analysed sperm length variation in terms of the intensity of sperm competition. This study investigates mating, sperm transfer and oviposition patterns in Drosophila bifurca, which possesses the longest sperm in the animal kingdom (about 6 cm). The prediction is that sperm gigantism should prevent male–male interaction. In this study, we examine how sperm transfer varies as males mate with a series of females, and how female receptivity changes with time after mating. As predicted, we found an extremely limited overlap of ejaculates owing to (1) reduced sperm transfer to females that had already mated, and (2) female remating depended both on the amount of sperm transferred and the modes of egg laying. The amount of sperm transferred to the female is discussed in relation to the peculiar morphology of the male reproductive tract and to sexual dimorphism and ecological hypotheses.     

Morphology of the female reproductive system of European pea crabs (Crustacea,Decapoda, Brachyura,Pinnotheridae)

Commensal pea crabs inhabiting bivalves have a high reproductive output due to the extension andfecundity of the ovary. We studied the underlying morphology of the female reproductive system in the Pinnotheridae Pinnotheres pisum, Pinnotheres pectunculi and Nepinnotheres pinnotheres using light microscopy and transmission electron microscopy (TEM). Eubrachyura have internal fertilization: the paired vaginas enlarge into storage structures, the spermathecae, which are connected to the ovaries by oviducts. Sperm is stored inside the spermathecae until the oocytes are mature. The oocytes are transported by oviducts into the spermathecae where fertilization takes place. In the investigated pinnotherids, the vagina is of the “concave pattern” (sensu Hartnoll 1968 ): musculature is attached alongside flexible parts of the vagina wall that controls the dimension of its lumen. The genital opening is closed by a muscular mobile operculum. The spermatheca can be divided into two distinct regions by function and morphology. The ventral part includes the connection with vagina and oviduct and is regarded as the zone where fertilization takes place. It is lined with cuticle except where the oviduct enters the spermatheca by the “holocrine transfer tissue.” At ovulation, the oocytes have to pass through this multilayered glandular epithelium performing holocrine secretion. The dorsal part of the spermatheca is considered as the main sperm storage area. It is lined by a highly secretory apocrine glandular epithelium. Thus, two different forms of secretion occur in the spermathecae of pinnotherids. The definite role of secretion in sperm storage and fertilization is not yet resolved, but it is notable that structure and function of spermathecal secretion are more complex in pinnotherids, and probably more efficient, than in other brachyuran crabs. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.