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.     

Atebbania Bernasconii, a New Genus and Species from Subterranean Waters of the Tiznit Plain, Southern Morocco (Gastropoda: Hydrobiidae)

The new genus Atebbania is described for a new hydrobiid species,A. bernasconii, which lives in groundwaters of southern Morocco.Of the Hydrobiidae genera with known shell and anatomy Moitessieriais the closest to Atebbania. The two share the following characters:shell elongate; teleoconch surface with evident microsculpture;seminal receptacle absent. Atebbania is distinguished from Moitessieriaby: shell ovate; penis with a) apical stylet, b) one lobe bentdownward on left (inner) side and c) penial duct running nearthe right (outer) side; posterior end of foot indented. Atebbaniabernasconii n. sp., lives in a limited area, the Tiznit plainin southern Morocco. Other stygobiont hydrobiids from otherareas of Morocco are currently being studied. None of them appearsto be closely related to A. bernasconii or to the species ofMoitessieria (Received 10 February 1998; accepted 30 April 1998)     

Dorymenia troncosoi sp. nov. (Mollusca Solenogastres: Proneomeniidae), a new species from the South Shetland Islands (Antarctica)

During the Spanish oceanographic expedition BENTART '95, carried out in Antarctic waters off Livingston Island (South Shetland Islands), five small specimens of a new species of Solenogastres-Cavibelonia, Dorymenia troncosoi sp. nov., were collected at a depth of 65–240 m on a silt bottom. The species is characterized by the presence of a pallial cavity with four pouches (a dorsal, two lateral and a ventral pouch), seminal receptacles that open into the spawning ducts through a short duct, radula having 9–11 teeth per row (1 central and 4–5 lateral, all the same size), the end of the copulatory spicules having a cross-section in the shape of a four-pointed star, and abdominal spicules present. These characteristics separate this species from other species of the genus, particularly from Dorymenia profunda, which is the most similar. Received: 9 April 1998 / Accepted: 12 June 1998     

Anatomy of Bathyacmaea secunda Okutani, Fujikura & Sasaki, 1993 (Patellogastropoda: Acmaeidae)

Anatomy of a vent-endemic patellogastropod limpet, Bathyacmaeasecunda, was examined by gross dissection and serial sections.It was revealed that B. secunda is anatomically distinct fromother patellogastropod taxa in that (1) the intestine runs throughthe ventricle; (2) the ventral approximator muscle of odontophoralcartilages is ventrally single-layered; (3) there is a pairof radular teeth with long shafts; and (4) the statocysts areisolated from pleural and pedal ganglia. In addition, the speciesis characterized by short pallial margin papillae, lack of pallialstreaks, presence of a ctenidium, obliquely tubular salivaryglands, simple gut configuration and acmaeoidean type of buccalmass musculature and odontophoral cartilages. The comparisonin anatomical and shell microstructural characters suggeststhat B. secunda shares no unique similarities with the Patelloideaor Neolepetopsidae and, therefore, is not closely related tothese groups. In contrast, a number of similarities were foundbetween B. secunda and acmaeid species. These results supportthe current systematic position of Bathyacmaea within the Acmaeidae,although anatomical data from some related genera are stillinsufficient. (Received 3 September 2005; accepted 25 January 2006)     

HOMOLOGY OF THE PALLIAL AND PULMONARY CAVITY OF GASTROPODS

The development and morphology of the pallial and pulmonarycavities of various gastropods has been investigated using epoxy-resinserial sections. In the veliger larvae of Cellana sandwicensis(Patellogastro-poda), Gibbula adansonii (Vetigastropoda), Modulustectum (Caenogastropoda) and Ovatella myosotis (Pulmonata) theformation of the pallial cavity is nearly identical. After shellformation a shallow dorsal pallial groove develops beneath themantle edge. During the late veliger stage, the ectoderm formsa deep invagination along the bottom of the pallial groove onthe right side of the larva, giving rise to the pallial cavity.In the ellobiid O. myosotis the pallial cavity becomes the lung(=pulmonary cavity), without any major post-metamorphic modification.Thus, the lung of this species is clearly homologous with thepallial cavity of prosobranchs. The lung of pulmonates withveliger development, as well as of fresh water basommatophoransand stylommatophorans, can be shown to be homologous by comparisonof adult morphology. In contrast to previous views, the pulmonatelung should be regarded as truly homologous with the pallialcavity of prosobranchs and opisthobranchs. In the onchidiidpulmonate Onchidium cf. branchiferum, the larval pallial cavityshifts posteriorly after metamorphosis, where it gives riseto a lung and a cloaca. Contrary to previous interpretations,it can be shown that the onchidiid lung is homologous with atleast part of the pallial cavity. Smeagol climoi has only asmall pallial cavity and no separate lung. The previously described‘lung’ is shown to be a gland. The re-evaluationof the development and morphology of the pulmonate lung hasimportant systematic implications: (1) The pulmonary cavitydoes not represent a synapomorphic character of pulmonates.(2) The gymnomorphs cannot be separated from the remaining pulmonatesbased on lung development. (3) The lack of a lung in the smeagolidsmight give reason to reconsider this group's systematic placementwithin the pulmonates.     

A hypothesis about the origin of sperm storage in the Eubrachyura, the effects of seminal receptacle structure on mating strategies and the evolution of crab diversity: How did a race to be first become a race to be last?

The origins and evolution of sperm storage in Brachyura are enigmatic: sperm is either stored in seminal receptacles, accessible via the vulvae on the sixth thoracic sternite, or in spermathecae at the border between the seventh and eighth sternites. Crabs with spermathecae are collectively referred to as “podotremes” while crabs with seminal receptacles belong to the Eubrachyura. The position of gonopores is the primary basis for subdividing the Eurachyura into the Heterotremata (female vulvae + males with coxal gonopores) and Thoracotremata (female vulvae + males with sternal gonopores). We present a hypothesis about the evolution of seminal receptacles in eubrachyuran female crabs and argue that the sternal gonopore has been internalized into chitin-lined seminal receptacles and the vulva is in fact a secondary aperture. The loss of some or all of the ancestral chitinous seminal receptacle lining was linked to ventral migration of the oviduct connection. Male and female strategies are to maximize gamete fertilization. The most important variable for females is sperm supply, enhanced by long-term storage made possible by the seminal receptacle. To maximize their fertilization rates males must adapt to the structure of the seminal receptacle to ensure that their sperm are close to the oviduct entrance. The major evolutionary impetus for female mating strategies was derived from the consequences of better sperm conservation and the structure of the seminal receptacle. The advantages were all to the females because their promiscuity and sperm storage allowed them to produce more genetically variable offspring, thereby enhancing variation upon which natural selection could act. We extend our arguments to Brachyura as a whole and offer a unifying explanation of the evolution of seminal receptacles, comparing them with the spermathecae found in “Podotremata”: they were independent solutions to the same problem: maintaining sperm supply during evolutionary carcinization.Explanation of eubrachyuran mating strategies requires analysis of the mating–moulting link, indeterminate vs. determinate growth format and seminal receptacle structure. Two alternatives for each of these characters means that there are eight possible outcomes. Six of these outcomes have been realized, which we term Portunoid, Majoid, Eriphoid, Xanthoid, Cancroid, and Grapsoid–Ocypodoid strategies, respectively. Mapping these characters on to a workable phylogeny (wherein some changes to the seminal receptacle + moulting–mating links are assumed to have occurred more than once) produces the following relationships: Portunoids + Majoids are a sister group to the rest of the Eubrachyura, which fall into two sister groups, Eriphoids + Xanthoids and Cancroids + Grapsoid–Ocypodoids and the “Podotremata” is sister group to all the Eubrachyura. We conclude that what began as a race to be the first to mate was turned on its head to become a race to be last, by the evolutionary changes to the seminal receptacle. Eubrachyuran females were advantaged by greater reproductive autonomy, more opportunity to mate with other males, resulting in more genetically variable progeny and leading to the evolution of much greater taxonomic diversity compared to “podotremes”.     

Functional anatomy of the female reproductive systems of two spider crabs (Decapoda,Majoidea)

To better understand the mating systems of majoid crabs, we studied the functional anatomy of the female reproductive systems of the spider crabs Leurocyclus tuberculosus and Libinia spinosa, comparing them with those of other Majoidea. Adult females were measured and dissected, and their reproductive systems described macroscopically and histologically. In females of both species, the seminal receptacles are paired globular structures of ecto‐mesodermal origin. The mesoderm‐derived region is lined by a stratified epithelium. The anchoring, proliferative, and secretory strata are clearly recognizable . The ectoderm‐derived region is lined by a simple cylindrical epithelium underlying a cuticle that increases in thickness toward the vagina. The transition between the ectoderm and mesoderm‐derived regions is abrupt, with differences between the studied species: Li. spinosa has a “velum,” whereas Le. tuberculosus presents prominent “folds.” In both species, the position in which the oviduct is connected to the seminal receptacles is intermediate between the dorsal and ventral types previously described in other eubrachyurans. The seminal receptacles of the studied species show four different conditions, which can be distinguished macroscopically based on their shape and amount of sperm stored. We compare our data with those from other Majoidea in an attempt to determine whether the morphology of the seminal receptacles is related to different mating strategies or behaviors.     

REPRODUCTIVE BIOLOGY OF THE SESSILE GASTROPOD VERMICULARIA SPIRATA (CERITHIOIDEA: TURRITELLIDAE)

The reproductive and developmental biology of the sessile gastropodVermicularia spirata (Philippi, 1836), collected from the FloridaKeys, was studied from living and preserved material. Individualsof this species attach themselves to a variety of substrata,but were mainly found embedded in the white sponge Geodia gibberosa.Pallial reproductive structures of both sexes of V. spiratawere found greatly to resemble those of Turritella communis,a free-living member of the same family. In both species, animalsof both sexes have open pallial ducts formed by two wide, recurvedlaminae. In the female of V. spirata, the laminae of the pallialoviduct serve as a capsule gland; a pair of side pockets representa fertilization pouch (possibly doubling as a copulatory bursa)and a seminal receptacle. The functional significance of theextensive, open, pallial sperm duct is not yet clear. Vermiculariaspirata is a protandrous hermaphrodite, and small males arefree-living; they become attached at about the time they undergosex reversal. Fertilization is probably brought about by malescrawling close to the apertures of the large, sessile femalesbefore releasing sperm. Egg capsules are brooded in the females'mantle cavities. Ova of about 300 µm diameter give riseto large (about 600 µm long) swimming-crawling veligerlarvae with shells of two and a half whorls. The veligers arecapable of metamorphosis at the time of hatching, but siblingsfrom one brood metamorphosed over a two-week period in the laboratory.Juvenile snails grew rapidly by filtering phytoplankton addedto their culture water. The life history of V. spirata is welladapted to assure fertilization and recruitment in a speciesotherwise committed to a sessile, filter-feeding existence. (Received 3 February 1989; accepted 19 April 1989)     

A new species of Acartia (Copepoda, Calanoida) from the Yellow Sea

A new species (Acartia hongi) of Acartia (family Acartiidae)predominating in the Korean coastal waters of the Yellow Seais described. This species closely resembles Acartia bifilosa,from which it can be easily distinguished by the co-occurrenceof the following morphological characters: the absence of rostralfilaments in both genders; the terminal spine on the femalefifth leg toothed only on the inner side; the first exopodalsegment of the male right fifth leg with a long seta; the distalsegment of the male left fifth leg with a rod-like appendage.Its distribution was discussed in comparison with two siblingspecies, Acartia omorii and Acartia hudsonica.     

MAINWARINGIA NEVILL, 1885, A LITTORINID GENUS FROM ASIATIC MANGROVE FORESTS, AND A CASE OF PROTANDROUS HERMAPHRODITISM

On the basis of reproductive anatomy and radula, the poorlyknown genus Mainwaringia Nevill, 1885, is shown to belong tothe Littorinidae. The genus is represented by two species, M.leithii (E.A. Smith, 1876) from India to Vietnam, and a newspecies, M. rhizophila, from Malaysia to Hong Kong. Both occurin mangrove forests. The reproductive anatomy of M, rhizophilais described in detail, and it is interpreted as a protandroushermaphrodite, a condition unique in the Littorinacea. In theintersexual and female stages both closed pallial oviduct andopen prostate are present in the mantle wall, suggesting thatthese structures are not strictly homologous. The affinitiesof Mainwaringiaare discussed, and it is concluded that it maybe most closely related to Littorina s.s. (Received 30 May 1986;     

Functional morphology of the female reproductive system of a crab with highly extensible seminal receptacles and extreme sperm storage capacity

We studied the functional morphology of the female reproductive system of the purple stone crab Danielethus crenulatus . The most remarkable feature is the relative storage capacity and extensibility of the seminal receptacles. These receptacles are a pair of simple sacs that lack internal structures dividing the internal lumen. Differences in seminal receptacle size and contents are accompanied by conspicuous changes in receptacle lining at a tissue level. Full seminal receptacles contain discrete sperm masses formed by hardened fluid and densely packed spermatophores. Different sperm masses are likely from different mates and their stratified disposition within the seminal receptacles is compatible with rival sperm displacement and last sperm precedence. Additionally, the anatomical structure of the vulva and vagina suggest active female control over copula. We discuss our results in the general context of sperm storage in brachyurans and the implications for the mating system of this species.     

THE ANATOMY OF CYLINDROTIS QUADRASI FORM OKINAWA ISLAND, JAPAN AND THE SUBFAMILIAL POSITION OF THE GENUS CYLINDROTIS MOLLENDORFF, 1895 (ARCHAEOPULMONATA: ELLOBIIDAE)

Cylindrotis quadrasi Mollendorff, 1895, the type species ofthe ellobiid genus Cylindrotis, was collected from Okinawa Island,southwest Japan. This is the first record of C. quadrasi inJapan and the third in the world. The anatomical charactersof this species, i.e. shell, head-foot, pallial cavity, radula,stomach, reproductive system, and central nervous system, aredescribed in detail for the first time. The significance ofthese observations is discussed and the subfamilial positionof Cylindrotis is reconsidered. Cylindrotis was included inthe Pythiinae, the most primitive group of the Ellobiidae, basedon the conchological characters alone. However, the male andfemale tracts of the reproductive system are entirely separated,and other internal organs exhibit highly derived conditions.Accordingly, Cylindrotis is transferred from the Pythiinae tothe Ellobiinae. Among the genera of the latter subfamily, Cylindrotisseems to be allied to Blauneria Shuttleworth, 1854 and LcucophytiaWinckworth, 1949. (Received 1 February 1993; accepted 7 June 1993)     

THE ANATOMY AND RELATIONSHIPS OF HYDROCOCCUS BRAZIERI (T. WOODS): Mollusca: Gastropoda: Rissoacea

Hydrococcus brazieri is an abundant, small gastropod which livesin brackish habitats on intertidal mud flats along the westernand southern coasts of Australia. It is typical of the Rissoaceain basic morphology but has a unique combination of charactersthat suggest it should be placed in a separate family, the HydrococcidaeThiele, 1928. The final assessment of the relationships of Hydrococcusmust, however, await more detailed information about relatedfamilies. The distinctive characters of Hydrococcus includea multispiral operculum with a central nucleus, a long, posteriorpallial tentacle with a short tentacle at its base, no metapodialtentacle, a penis bearing two accessory lobes, a bursa copulatrixwhich opens directly into the mantle cavity independently ofthe oviduct opening and rather long connectives between boththe pleural and oesophageal ganglia. There is no ctenidium butfolds on the pallial roof probably have a respiratory function. Hydrococcus graniformis Thiele and Ampullarina minuta T. Woodsare shown to be synonyms of Rissoina brazieri T. Woods. A clinebetween the forms typical of brazieri and graniformis is demonstrated. (Received 20 November 1980;     

HETEROSTOPHIC SHELLS AND PELAGIC DEVOLOPMENT IN TROCHOIDEANS: IMPLICATIONS FOR CLASSIFICATION, PHYLOGENY AND PALAEOECOLOGY

Four species of archeogastropods, presumed members of threesubfamilies of the trochidae, exhibit significant differencesin developmental modes and shell coiling. All four species havelecithotrophic development which is reflected in their inflatedpausispiral protoconchis; however, Margarites marginatus andLirularia succincta have benthic development in gelatinous masses,while Margarites pupillus and Calliostoma ligatum have pelagicembryos and swimming larvae with a potential for dispersal overa period of a week or longer. These modes cannot be deducedfrom the size of the egg, the size or shape of the protoconch,or the size or relative prominence of female pallial reproductivestructures. The protoconch of C. ligatum is orthostrophicallycoiled, but the protoconchs of the other three species are hyperstrophicallycoiled although their teleconchs are orthostrophic. These threetrochoidean species thus share with architectonicoideans, pyramidelloideans,opisthobranchs, and pulmonates the distinctive shell characterof heterostrophy, previously unreported for archaeogastropods. These observations, considered together with others reportedin the literature, show: (1) that developmental mode (pelagicor benthic) cannot be inferred from protoconch appearance ortaxonomy in major trochoidean ganera; (2) that significant dispersalpotential is present in the histories of some trochoidean archeogastropoids;and (3) that character sets (pallial reproductive structures,pairing during spawning, heterostrophic shell coiling) thoughtnot to occur below the mesogastropod level are found in theTrochoidea. These conclusions bring into question the usefulnessof these characters in defining higher gastropod taxa and raiseadditional questions concerning the ancestry of the higher gastropods. (Received 16 February 1989; accepted 25 June 1989)     

New species, anatomy, and systematic position of the hydrothermal vent and hydrocarbon seep gastropod family Provannidae fam.n. (Caenogastropoda)

The anatomy of the hydrothermal vent gastropod Provanna is described. It is shown to be a primitive neotaenioglossan with mainly plesiomorphic features in all systems, including open pallial gonoducts, epiathroid nervous system with accessory pedal ganglia and a circulatory system where a considerable portion of the venous blood from the coelomic spaces enters the pallial skirt. Apomorphic features include the possession of an anterior notch, or short canal in the shell (and mantle edge), an annulated sensory pallial tentacle, paraspermatozoa modificd as nurse cells. multiple seminal receptacles, a posterior bursa copulatrix, and a renal oviduct with three distinct regions. the middle one expanded. Comparisons are madc with Abyssochrysos, a genus assumed to be a relict member of the Loxonematoidea, which appears to be the closest known relative. A new family. Provannidae fam.n., is created and tentatively included in the Loxonematoidea. The following new specics are described: Provanna segonzaci sp.n. (from hydrothermal vents at the Fiji Back Arc), P. laevis sp.n. (from hydrothermal seeps in the Gulf of California), P. sculpta sp.n. and P. admetoides sp.n. (from hydrocarbon seeps in the Gulf of Mexico). New records are added for previously described species of Provanna.