Ultrastructure and functional morphology of male genital organs and spermatophore formation inProtodrilus (Polychaeta,Annelida)

Summary The structure and functional morphology of lateral organs and sperm ducts, as well as the mechanisms of spermatophore formation and transfer, are investigated by means of light and electron microscopy in the genusProtodrilus. The sperm ducts are simple, ciliated, intercellular gonoducts with a funnel section surrounded by a thin muscle layer and a tube section opening externally in the anterior region of the lateral organs. No glands are present in the sperm ducts. The lateral organs are formed by long epidermal invaginations enclosing an elongate lumen into which numerous cilia project and a large number of glands open. Five to ten different gland types with strikingly distinctive secretory granules are found in the different species. In addition, special supporting cells, the so-called sponge cells, sensory cells and an underlying nervous tissue are developed in the lateral organs. It is stated that apart from some similarities to the ventral atrium ofNerilla antennata no corresponding organs are known within the Annelida. It is argued that inProtodrilus the spermatophores are formed by the lateral organs as there are a high number of glands opening into the lumen of the organ. The possible origin and genesis of the male gonoducts as well as the mode of spermatophore transfer inProtodrilus is discussed.Abbreviations used in the figures bl basal lamina - cc coelomic cell - ci ciliated cell - cir ciliary root - cr ciliary ring - cu cuticle - cv bs contractile ventral blood sinus - d dissepiment/septum - dbs dorsal blood sinus - es euspermatozoa - f funnel - fi filament - g gut - glo gland openings - lgl lateral organ gland - lm longitudinal muscle - lo lateral organ - lu lumen - mi mitochondrion - mt microtubules - mu muscle - mv microvilli - mvc microvillar crown - n nucleus - ne nervous tissue - o opening - ps paraspermatozoa - rer rough endoplasmatic reticulum - s spermatozoa - sc sponge cell - sg salivary gland - spd sperm duct - spdo sperm duct opening - t tube - tm transverse muscle - vc ventral ciliary band     

Functional morphology of the mammiliform penial glands ofLittorina saxatilis (Gastropoda)

Summary The functional morphology of the mammiliform penial glands ofLittorina saxatilis has been investigated with both light and electron microscopy. These penial glands line the ventral edge of the penis and orient with the female mantle during copulation. Secretions are released from the penial glands to this interface where they probably function in adhesion. The penial gland secretions comprise heterogeneous granules as well as apocrine and mucous secretions. The heterogeneous granules are produced in separate multicellular glands arranged in a series of lobes that lie outside a thick smooth muscle layer enclosing the lumen. Each glandular lobe is surrounded by a thin layer of smooth muscle. Secretions are transported in individual cellular processes that pass through the thick smooth muscle layer and empty into the lumen. Surrounding the lumen is an epithelium containing apocrine secretory cells as well as occasional goblet-type, mucous cells. The combined action of the muscles forces secretions out of the lumen through the penial papilla, onto the external surface of the mammiliform penial gland. Longitudinal muscles extend into the penial papilla enabling its protrusion or retraction. Retraction of the penial papilla following secretion release is thought to create negative pressure beneath the penial gland producing suction adhesion. The visco-elastic properties of the penial gland secretion are qualitatively different from foot mucus and may represent specialization to an adhesive function.     

Ultrastructure of copulatory organs in Turbellaria

Summary The copulatory organs in Macrostomum sp. and Microstomum sp. contain simple tubular stylets which are intracellular specializations. The stylet in Macrostomum sp. is produced in a syncytium covering part of the prostatic vesicle. The proximal region of the stylet surrounds the vesicle which contains six prostatic gland ducts and six accessory (sensory) cells containing ciliary rootlets. The stylet in Microstomum sp. is produced in an extension of a syncytium which lines the combined seminal-prostatic vesicle. The stylet is connected to the combined vesicle by a narrow bridge of matrix syncytium through which sperm, prostatic gland products and sensory cilia pass from the vesicle to the stylet lumen. In both species the matrix syncytium can be interpreted as a specialized terminal end of the male canal epithelium. Stylets of Turbellaria and other lower Metazoa are discussed in regards to structure (one or several pieces) and location (in separate cells, in a syncytium, or extracellular).Abbreviations used in figures ac accessory cell - b basal body - c cilium - cv combined vesicle - d prostatic gland duct - dc degenerative cell - di dictyosome - e epidermis - ed ejaculatory duct - g prostatic gland cell - h hemidesmosome - i intercellular matrix - im internal muscle - in intestine; - l lumen of male canal - lm longitudinal muscle - m matrix syncytium - mc male canal epithelial cell - mi microfilaments - mt microtubules - mu muscle cell - mv microvilli - n nucleus - np nerve process - ns neurosecretory (?) granule - p prostatic vesicle - pv prostatic part of combined vesicle - r rootlet - s stylet - sm stylet material - sp sperm - sv seminal part of combined vesicle     

Ultrastructure of the copulatory organ of Haplopharynx quadristimulus and its phylogenetic significance (Plathelminthes,Haplopharyngida)

Summary The male reproductive system in Haplopharynx quadristimulus consists of paired testes, sperm ducts, seminal vesicles, seminal ducts, a copulatory organ containing prostatic vesicle and stylet apparatus, and the male canal. By electron microscopy all components appeared to be regional specializations of a canal extending from the testes to the body wall and lined by a multiciliated epithelium. The epithelium of the stylet apparatus contained six different cell types. One cell type (matrix syncytium) formed the stylet and the other five were located distal to the stylet/prostatic-vesicle junction along the male system epithelium. Each cell type was attached to the supporting intercellular matrix at a different level along the stylet apparatus. All cell types extended to the distal end of the stylet apparatus regardless of where they originated along its length. The cells in the apparatus lacked cilia, but one of the cell types contained rootlets. Modified rootlets or rootlet derivatives were possibly present in another cell type in the form of rootlet-like ribbons. The findings support the monophyly of the Macrostomida Haplopharyngida (by common occurrence of a matrix syncytium) and at the same time suggest their separation as two distinct taxa (by differences in the structure of the prostatic vesicle and other parts of the stylet apparatus).Abbreviations a accessory spine - c circular muscle - ce centriole - ci cilium - di dictyosome - e epithelial cell - ed ejaculatory duct - ep epidermal cell - f rootlet-like ribbon - g prostatic gland cell neck - g1 type I gland cell granules - g2 type II gland cell granules - g3 type III gland cell granules - h hemidesmosome - i intercellular matrix - im internal muscle - j septate junction - l stylet apparatus lumen - le spine lateral extension - lm longitudinal muscle - m matrix syncytium - mc male-canal epithelial cell - me male canal - mp male pore - mt microtubules - mv microvilli - n nucleus - nc nerve cell body - np nerve process - om oblique muscle - p prostatic vesicle epithelial cell - pv prostatic vesicle - r rootlet - s stylet - sa stylet apparatus - sc sensory receptor - sd sperm duct - se seminal duct - sl stylet lumen - sp spot desmosome - sr sperm - sv seminal vesicle - t terminal web - te testis - u ultrarhabdite - z zonula adhaerens - 2 cell type 2 - 3 cell type 3 - 4 cell type 4 - 6 cell type 6     

Reconstruction of the musculature of <Emphasis Type="Italic">Magelona</Emphasis> cf. <Emphasis Type="Italic">mirabilis</Emphasis> (Magelonidae) and <Emphasis Type="Italic">Prionospio cirrifera</Emphasis> (Spionidae) (Polychaeta,Annelida) by phalloidin labeling and cLSM

Recent investigations have suggested that a lack of circular muscle fibers may be a common situation rather than a rare exception in polychaetes. As part of a comparative survey of polychaete muscle systems, the F-actin musculature subset of Magelona cf. mirabilis and Prionospio cirrifera were labeled with phalloidin and three-dimensionally analyzed and reconstructed by means of cLSM. Obvious similarities are sublongitudinal lateral, circumbuccal, palp retractor, dominating dorsal longitudinal, perpendicular lateral and ventral transverse muscles. Differences between M. cf. mirabilis and P. cirrifera are: (1) two types of prostomial muscles (transversal and longitudinal) in M. cf. mirabilis versus one type (diagonal) in P. cirrifera; (2) one type of palp muscles (longitudinal) in M. cf. mirabilis versus three types (longitudinal, diagonal, circular) in P. cirrifera; (3) five ventral longitudinal muscles (ventromedian, paramedian, ventral) in M. cf. mirabilis versus four (two paramedian, two ventral) in P. cirrifera. Ventral and lateral transverse fibers are present in the thorax, but absent in the abdomen of M. cf. mirabilis. The triangular lumen of the pharynx in M. cf. mirabilis is surrounded by radial muscle fibers; three sets of pharynx diductors attach to its dorsal side. The unique features of P. cirrifera are one pair of brain muscles and segmentally arranged dorsal transverse muscles, the latter located outside the longitudinal muscles. The transverse lateral muscles are restricted to the sides and lie beneath the longitudinal muscles, a pattern described here for the first time. A true, outer layer of circular fibers is absent in both species of Spionida that were investigated.     

Comparative morphology, homologies, and functions of the male system in oribatid mites (Arachnida: Acari)

The penis is basically a double-walled oval cone. The weak type penis (only 2 species) has a weak tongue, lacks a bridge, and is elongate. The strong type (all others) has a bridge, a strong tongue, thicker walls, and is short. An accessory gland and a common vas deferens always open into the inner cup lumen (= ejaculatory duct). The massive tongue muscles may open the penal orifice. A pair of penal retractor muscles originate on the body wall. Penal protrusion and perhaps partial extrusion of stalk substance is by hemolymph pressure. The penis is completely homologous to the ovipositor. The genital discs are cuticular cups containing glandular tissue and are retractible by muscles originating on the body wall. The minute, rod-like, immobile sperm are mixed with seminal fluid and stalk material secreted by seminal vesicle cells. This mixture is carried via the vase deferentia by peristalsis to the penis. Semen and stalk substance (protein) are somehow separated in the ejaculatory duct into separate pools, with stalk substance nearest the penal orifice. Upon penal protrusion, a bit of stalk material is extruded and fastened to the ground, and upon raising of the mite's body the stalk is “drawn out.” Finally, the ball of semen, adhering to the stalk tip, is pulled through the penal orifice.     

Unusual granules in the ejaculatory duct of a Microphthalmus species (Polychaeta,Annelida)

Summary The paired prominent ejaculatory ducts of the hermaphroditic polychaete Microphthalmus cf. listensis are surrounded by gland cells the processes of which penetrate the ducts themselves. These cells produce, in separate regions, two different types of spherical granules. Type I is composed of an electron dense and an electron lucent part. Type II granules contain a tubular filament that forms a single or double spiral in the periphery of a more or less unstructured electron dense material. Golgi vesicles give rise to this granule type. During the passage of sperm, these granules are obviously discharged into the lumen of the duct. Here they change form and probably dissolve. Their function is as yet unknown; capacitation of sperm is assumed.     

Morpho-functional variety of the coxal glands in cheyletoid mites (Prostigmata). II. Cheyletidae

Trombidiform mites are characterized by the presence of several paired glands in the anterior body portion united by a common conducting duct (podocephalic canal). Apart from the acinous (salivary) glands the podocephalic system includes a pair of tubular coxal glands (CGs) responsible for osmoregulation. The aim of the present study was to figure out how functional changes of acinous glands reflect on the corresponding CG. For this purpose, the anatomy and fine structure of the CG were analyzed in two mite species, Bakericheyla chanayi and Ornithocheyletia sp. (Cheyletidae), which have a different composition of their single acinous gland.The results showed that in both species the CG lacks a filtering saccule. It is composed of the proximal and distal tubes and leads into a cuticle-lined excretory duct. Both tubes demonstrate a similar species-specific fine structure. They are characterized by an extensive system of apical membrane invaginations (internal canals) associated with numerous large mitochondria. Local areas of modified internal canals were regularly observed in both species. They contain structures resembling those constituting filtering slit diaphragms of other animals.In O. sp., CG cells in addition demonstrate features characteristic of protein-like secretion. Apparently this correlates with the loss of true salivary glands in this species, as its acinous gland was previously assumed as silk producing. Contrary to this, the CG of B. chanayi shows no kind of granulation, which coincides with the presence of a salivary portion in its complex acinous gland.The microtubule-rich intercalary cells at the base of the excretory duct were associated with special muscles presumably regulating the dilation of the duct lumen. These cells might represent a basic feature common to different types of podocephalic glands.     

Ultrastructure of the Male Accessory Glands and Sperm Ducts of Cylindrotaenia hickmani(Cestoda,Cyclophyllidea)

Abstract The ultrastructure of unicellular accessory glands (= prostate glands) and external male ducts of the cestode Cylindrotaenia hickmaniare described. Accessory glands open into the lumen of the external common sperm duct (= external vas deferens). The gland cells contain abundant endoplasmic reticulum, Golgi bodies and secretory bodies, and have elongate necks that pierce the apical cytoplasm of the duct. Cell contact with the apical cytoplasm of the sperm duct is mediated by septate desmosomes. Accessory glands secrete spherical particles, with a diameter of approximately 70 nm, that adhere to spermatozoa. The roles of these accessory glands may relate to activity of the sperm or development of the female system after insemination. Paired sperm ducts arise from testes, and unite to form a common sperm duct. Each duct consists of a tubular anucleate cytoplasmic region which is supported by nucleated cytons that lie sunken in the parenchyma. The apical cytoplasm of the paired sperm ducts (= vasa efferentia) possesses apical microvilli and abundant mitochondria, but few other cytoplasmic features. The apical cytoplasm of the common sperm duct possesses sparse apical microvilli and numerous electronlucent vesicles. The male gonoducts form an elongate syncytium which is markedly polarized along the length of the ducts. The ducts also display apical–basal polarity in that sunken nucleated cytons support the apical cytoplasm which in turn has distinct basal and apical domains.     

Morphological and histological study of the genital ducts ofCryptazeca monodonta (Pulmonata,Orthurethra), with special emphasis on the auxiliary copulatory organ

Summary The genital system ofCryptazeca monodonta is very similar to those reported for semidiaulic stylommatophores, but with some specific features. The fertilization pouch is simple and surrounded by subepithelial goblet gland cells. The spermoviduct has two different grooves: the oviductal channel and the spermatic groove, which run together with a blind-ending allospermiduct that opens into the lumen of the free oviduct. The vagina has a thick muscular wall with numerous pigmentary cells embedded in it. The vas deferens diverges from the spermiduct and becomes mainly glandular just before it joins the penis. This genital system is equipped with an auxiliary copulatory structure that consists of two independent and complementary organs. One of them is located just before the fusion of the free oviduct and the spermatheca stalk lumina and consists mainly of a thick mass of connective tissue. The other is a muscular sarcobellum located inside the penis. Both these organs are covered by small papillae whose connective cells are stacked. Each papilla has a solid spine on its top, which seems to be of mesenchymatous origin. As in other stylommatophores, the auxiliary copulatory organ is equipped with an adjoining gland, which inCryptazeca is next to the sarcobellum.Abbreviations ACO auxiliary copulatory organ - ag albumen gland - al allospermiduct - c connective aggregate of ACO - cc connective cells of ACO papillae - cf connective fibres - ep epiphalus - fo free oviduct - fp fertilization pouch - gl.c gland cells of sarcobellum - hd hermaphroditic duct (distal portion) - m muscle fibres - ov oviduct - p penis - pc penial caecum - pp papillae of ACO - pr prostatic gland - prm penial retractor muscle - p.sh penial sheath - s spermatheca - sc sarcobellum - SEM scanning electron micrograph - sov spermoviduct - sp spine - spd spermiduct - ss spermatheca stalk - v vagina - vd vas deferens     

The morphogenesis of spermathecae and spermathecal glands in Drosophila melanogaster

Sperm storage in female insects is important for reproductive success and sperm competition. In Drosophila melanogaster females, sperm viability during storage is dependent upon secretions produced by spermathecae and parovaria. Class III dermal glands are present in both structures. Spermathecal glands are initially comprised of a three-cell unit that is refined to a single secretory cell in the adult. It encapsulates an end-apparatus joining to a cuticular duct passing secretions to the spermathecal lumen. We have examined spermatheca morphogenesis using DIC and fluorescence microscopy. In agreement with a recent study, cell division ceases by 36 h after puparium formation (APF). Immunostaining of the plasma membrane at this stage demonstrates that gland cells wrap around the developing end-apparatus and each other. By 48–60 h APF, the secretory cell exhibits characteristic adult morphology of an enlarged nucleus and extracellular reservoir. A novel finding is the presence of an extracellular reservoir in the basal support cell that is continuous with the secretory cell reservoir. Some indication of early spermathecal gland formation is evident in the division of enlarged cells lying adjacent to the spermathecal lumen at 18 h APF and in cellular processes that bind clusters of cells between 24 and 30 h APF.     

Anatomy and ultrastructure of ventral pharyngeal organs and their phylogenetic importance in Polychaeta (Annelida)

Summary A comparative anatomical and ultrastructural study of ventral pharyngeal organs (pharyngeal bulbs) was carried out in two species of the Dinophilidae: Dinophilus gyrociliatus and Trilobodrilus axi. Special attention was paid to the fine structure of the stomodeal epithelium, cuticle, glands, muscles, and myoepithelial junctions. The differences between the species are very slight. The pharyngeal organ of the Dinophilidae is characterized by the following features: solid muscle bulbus made up of muscle cells only, bulbus muscle cells with two myofilament systems crossing at an angle of about 90°, gap junctions between these muscle cells, bulbus projects into a pharyngeal sac and bears rostrally a specific epithelium and cuticle, no bulbus glands, no investing (= sagittal) muscles, specific cuticle ultrastructure, cilia of ascending oesophagus with asymmetric tips, specific structure and position of salivary gland openings. The phylogenetic importance of these structures is discussed. Some of these characters are clearly autapomorphic features of the Dinophilidae and no common derived structures to other families with a ventral pharyngeal organ are present. Therefore, it is most likely that the dinophilid pharyngeal organ evolved independently. These findings do not agree with the hypothesis of the unity of the archiannelid families (Polygordiidae, Protodrilidae, Saccocirridae, Nerillidae, Dinophilidae, and Diurodrilidae) established on the basis of an assumed structural similarity of their ventral pharyngeal organs.Abbreviations bb basal body - bep bulbus epithelium - bl basal lamina - bm bulbus muscle - c cilium - cc coelenchyme cell - cm circular muscle - cr caudal rootlet - cu cuticle - dblm dorsal bulbus longitudinal muscle - dlm dorsal longitudinal muscle - dsn dorsal stomatogastric nerve - dy dyad - el electron-dense layer - fl fibrous layer - fi filaments - g Golgi apparatus - gl gland cell - hv homogeneous vesicle - l lipid droplet - la external lamina - lal lamellar layer - ll lower lip - lm longitudinal muscle - ly lysosome - m mitochondrion - mo mouth opening - mt microtubule - mv microvillus - mvp microvillar process - n nucleus - nu nucleolus - oes oesophagus - pcom preoral commissure - phf pharyngeal fold - phl pharyngeal lumen - phs pharyngeal sac - pms peripheral myofilament system - r rootletlike structure - rer rough endoplasmic reticulum - rr rostral rootlet - s sarcoplasmic reticulum - sc salivary canal - scom suboesophageal commissure - sd septate desmosome - ser smooth endoplasmic reticulum - sg secretory granule - sgl salivary gland - sn stomatogastric nerve - st stomach - step stomodeal epithelium - tep transitional epithelium - tf tonofilaments - va vacuole - vlm ventral longitudinal muscle - vsn ventral stomatogastric nerve - z z-element - za zonula adherens     

Internal fertilization in Lepidogalaxias salamandroides Mees (Pisces: Lepidogalaxiidae)

The gonads of Lepidogalaxias salamandroides are structurally distinctive. A duct, the surface of which is ciliated, is located on the ventral surface of each ovary and may be a modification of the ovarian tunica. The testes are small and the majority of the testicular structure functions as a sperm storage area. Secondary requisite sex characters are present in the male and the function of a modified scale sheath in copulation is unique among teleosts. Fertilization is internal and a viscous mucus mass secreted during copulation may serve as a copulatory plug. It is postulated that internal fertilization within this species has arisen as a response to sperm competition.     

The fine structure of Gnathostomulid reproductive organs

Summary The male copulatory organs of five species of Gnathostomulida Scleroperalia have been studied by TEM techniques. These observations provide a more solid basis for classification in the light microscope: inLabidognathia longicollis (fam. Mesognathariidae) the stylet is composed of eight, and inSemaeognathia sterreri, Gnathostomula jenneri, Gnathostomula mediterranea andGnathostomula microstyla (Gnathostomulidae) of ten stylet rods. Each rod consists of a microtubule-filled inner rod, and of an outer rod, filled with crystallized inclusions. The inner rods are continuous with eight — or ten — rod formation cells which are located in the proximal stylet sack. Bipartition of rods occurs by a longitudinal invagination of the basement lamina, underlying the rod cells and the gland cells and continuous with that of the body wall epithelium. InLabidognathia, the outer rods are interlocked, in Gnathostomulidae, the stylet rods are surrounded by an extracellular (cuticular) tube-like stylet sheath of variable fine structure, which is believed to provide extra rigidity. In the species investigated, one single stylet gland, consisting of a monolayered epithelium showing different gland cell types, surrounds the stylet. In the apical gland cell portions, medially and distally membrane-bound secretory granules lie adjacent to the stylet sheath. In Gnathostomulidae, two anterior gland cells are seen in connection with the formation of the stylet sheath. In the muscular sheath the cross-striated fibers, basically derived from the longitudinal body wall musculature, show a tendency towards helical and circumferential arrangement. Musculature is especially prominent in the proximal stylet sack, which is rather a propulsive element than a sperm-storing vesicle, and lacks glands. InGnathostomula species, atrial cells underlie the distal tip of the stylet. The entrance into the male opening is lined with ciliary receptor cells and specialized gland cells.Stylet evolution in Scleroperalia is characterized by progressive differentiation of the muscular sheath, in particular of the proximal stylet sack, and of the stylet — the occurrence of a stylet sheath is seen in connection with increasing diversity of stylet shape.Abbreviations ac atrial cell(s) - ag anterior gland cell(s) - b bursa - bl basal lamina - c rod-crystal in outer rod - cj cuticle of jaw - d desmosome - di dictyosome - e body wall epithelium - ej pharyngeal epithelium - g stylet gland (cell) - gm median gland cell - i gut (cell) - ir inner rod - jc junctional complex - m muscular layer - mo male opening - mv microvillar protrusions - nu nucleus - o ovary - or outer rod - po proximal opening of the proximal stylet sack - ps proximal stylet sack - r stylet rod - rc rod cell - sg secretory granule - sj septate junction - sp sperm - ss stylet sheath - st stylet - te testes - v ventral - z centriole     

Ultrastructure of the male reproductive organs in the interstitial annelid Sphaerosyllis hermaphrodita (”Polychaeta”, Syllidae)

 The reproductive organs of the simultaneous hermaphrodite Sphaerosyllis hermaphrodita (Syllidae, Exogoninae) were examined by TEM and reconstructed from ultrathin serial sections. Oocytes are produced in the 11–13th chaetigerous segments and then attached to the outer body surface. The male organs comprise a seminal vesicle, testes, sperm ducts and copulatory chaetae. The unpaired seminal vesicle is an uncompartmented cavity above the gut and within the chaetigerous segments 8–10. Its interior is lined with a layer of gland cells that degenerate as spermatogenesis in the vesicle proceeds. The testes are situated ventrolaterally, close to the seminal vesicle in the 9th chaetigerous segment. They contain cells at early stages of spermatogenesis, which are connected to one another by zonulae collares. The testes and seminal vesicle are enclosed in epithelia. Paired sperm ducts run ventrally from about the midline of the body under the seminal vesicle and into the parapodia of the 9th chaetigerous segment. There they open, together with the protonephridia of this segment, to the outside next to the stout copulatory chaeta. Each sperm duct consists of six cells, the luminal surface of which bears microvilli but no cilia. Only in animals with fully differentiated sperm does the small opening of the proximal duct cell in each duct give access to the seminal vesicle. The mode of sperm transfer is discussed. Accepted: 9 December 1996     

Données histologiques sur les fosses nasales et leurs annexes chezCrocodylus niloticus Laurenti etCaiman crocodilus (Linnaeus) (Reptilia,Crocodylidae)

Summary The vestibulum is very short and lined by stratified squamous epithelium which contains many alveolar cells. The cavum nasi proprium is exceedingly complex, with three conchal formations and a series of six recesses and sinuses. Olfactory epithelium lines the whole dorsal or dorso-medial half of the cavum, but not the deep sinuses. Non sensory respiratory epithelium lines the ventral or ventro-lateral half of the cavum, all the caviconchal recess, the posterolateral recess, the postturbinal sinus, the postconchal cavity, and the nasopharyngeal duct. In olfactory epithelium the proportion of sensory cells is about 61 % inCrocodylus and 59 % inCaiman; the ratio of sensory cells to supporting cells is about 2.6/1 in the former and 1.8/1 in the latter. Bowman's glands are sero-mucous and normally developed. As in other reptiles, the respiratory epithelium is composed with mucous and ciliated cells; but, in ventrolateral part of the cavum, there are also sero-mucous cells forming small multicellular glands. The hypertrophied lachrymal duct constitutes a very large naso-lachrymal not previously described gland. The lumen is lined by mucous and ciliated cells, the collet of each branched tubular gland by mucous cells and the glandular tubes by sero-mucous cells. Adult crocodilians lack a Jacobson's organ and there are no vomeronasal sensory epithelium in the cavum.In reptiles, aquatic way of life generally involves regression in olfactory epithelium, while Jacobson's organ (or, in Testudines, vomeronasal epithelium) persists and occasionally increases. In crocodilians, things are exactly reverse. After comparing with other Tetrapods, it seems likely that Jacobson's organ has been lost by terrestrial ancestor of crocodilians and birds. Now, only crocodilians posses olfactory epithelium, and naso-lachrymal gland gives them a supplementary protection, necessary in semiaquatic environment.     

Developmental sequence for the copulatory organs of Kontikia mexicana with remarks on taxonomic significance (Turbellaria: Tricladida: Geoplanidae)

Five specimens, presumably representing different developmental stages of the land planarian Kontikia mexicana (Hyman, 1939), were used to reconstruct the development of the copulatory apparatus in this species. The results support the notion that Kontikia differs from the closely related Caenoplana in its possession of a penis papilla. In the earliest stage available, a penis papilla was absent and other components were not differentiated. In a late-juvenile condition, the gonopore, seminal vesicle, and ejaculatory duct were present. The short penis papilla appeared to arise in this stage by elongation of the terminal tissue around the ejaculatory duct and its separation from the antral wall. The female canal was guarded by an epithelial fold and the glandular duct was present. In the mature condition, the penis papilla was more elongate, and the secretory (prostatic) region of the ejaculatory duct was functional. The female canal, guarded by an epithelial fold, was well-developed with enlarged glandular duct but lacking the posterior diverticulum and the sperm storage system associated with the ovovitelline ducts known in Kontikia orana Froehlich, 1955.     

A new viviparous acoel Childia vivipara sp. nov. with observations on the developing embryos, sperm ultrastructure, body wall and stylet musculatures

A free‐living viviparous acoel, Childia vivipara sp. nov., from the Gullmar fjord of the Swedish coast is described. The new species is assigned to the taxon Childia based on histological, ultrastructural and molecular sequence similarities. All available molecular markers (18S rRNA, 28S rRNA and histone H3) and several morphological characters, obtained using transmission electron microscopy and confocal scanning laser microscopy of whole mount specimen stained with TRITC‐labelled phalloidin, support the placement of C. vivipara in the taxon Childia. Childia vivipara and other Childia species share the following morphological synapomorphies: well‐developed copulatory organs built of tightly packed stylet needles, proximal part of the stylet inserted into the seminal vesicle, reversed body‐wall musculature, absence of ventral diagonal muscles, presence of dorsal diagonal muscles, and presence of ventral straight longitudinal muscles between frontal pore and mouth, 9 + 1 sperm axoneme structure, six distal sperm cytoplasmic microtubules, and extensive overlap of axonemes and nucleus. The new species can be easily distinguished from other Childia species by its viviparous mode of reproduction and single curved stylet. Observations on late embryonic development based on the oldest developing embryos are discussed.     

Structure,function, and evolutionary significance of the reproductive system in males of Hebrus ruficeps and H. pusillus (heteroptera,gerromorpha, hebridae)

In Holland, bugs of the species Hebrus pusillus and H. ruficeps have one generation per year and overwinter as unmated adults. Males have two testes with two follicles + vasa efferentia each, paired vasa deferentia and seminal vesicles, an ejaculatory duct, and a protrusible phallus comprising an articulatory apparatus, phallotheca, endosoma, and paired claspers. The skeletomusculature of this system is described (it has 12 paired and four unpaired muscles) and its functions in generating and transferring sperm (summarized in Figs. 70–75) are reconstructed from study of living bugs, dissections, whole mounts, and serial sections. Males of both species produce sperm >2 mm long from stem spermatogonia in the germarium of each follicle. Initial definitive spermatogonia divide synchronously three times to form clones of eight, interconnected, primary spermatocytes. These enlarge up to 43-fold in males of H. pusillus and 78-fold in those of H. ruficeps, undergo meiosis, and, after adult emergence, complete their differentiation into bundles of 32 sperm which coil transversely about the periphery of each follicle at its base. These begin to enter the vasa efferentia in mid August, rupture, and release their sperm into the seminal vesicles where they are stored overwinter. Most spermatocyte and spermatid cysts remaining in the testes degenerate in fall, leaving only stem spermatogonia and a few early spermatocysts in the germaria. Males of H. pusillus begin to mate the first warm days of spring but only the most persistent succeed. A male jumps on the back of a female, induces her to lower her ovipositor, and, within 12 min (@ 18–24°C), introduces the endosoma of his phallus up its shaft and fills his seminal duct with sperm. The female draws this into her gynatrial sac at the end of copulation and transfers it into her spermatheca in about 30 min, the sperm reversing themselves within it so that their heads face towards its mouth. The male may stay on her back for up to 2 hours and may copulate again up to three times before leaving to mate with other females. Males of H. pusillus may be sexually active for months after overwintering, because spermatogonia in their germaria reactivate in spring to produce additional sperm. Those of H. ruficeps do not and males mate successfully only until their supply of overwintered sperm is exhausted. The chromosome complement of H. pusillus males is 2N = 22 + XY. The X and Y chromosomes are of unequal length, form a pseudo pair at metaphase I, and segregate to opposite poles at anaphase I—the first instance of pre-reductional segregation of sex chromosomes to be recorded in the Gerromorpha. The chromosome complement of H. ruficeps may be 2N = 24 + XO but the nature of two chromosomes was not resolved. The single X segregates to half the spermatids at anaphase II.     

Morphological evidence for limited sperm production in the enigmatic Tasmanian cave spider Hickmania troglodytes (Austrochilidae,Araneae)

The male reproductive systems of spiders are highly diverse in structure across all major spider taxa. Little is known about this organ system in basal araneomorph spiders, especially Austrochiloidea; such knowledge is necessary for a more complete understanding of the evolutionary morphology of the male reproductive system in spiders. In the present study, we describe the male reproductive system of an austrochilid spider, the enigmatic troglophilic Tasmanian cave spider Hickmania troglodytes, using light and electron microscopic techniques. The male reproductive system consists of tubular testes leading into convoluted deferent ducts, which are fused close to the genital opening to an unpaired ejaculatory duct. Spermatogenesis occurs only in the subadult testes, whereas adult testes showed neither spermatogenic stages nor any generative tissue in all investigated specimens. The testes of adult males are drastically reduced in size compared with those of subadult males, but the deferent ducts are filled with large numbers of mature spermatozoa. Thus, our data suggest that males of H. troglodytes are sperm‐limited, but not necessarily sperm‐depleted as described for certain orb‐weaving spiders. Due to the absence of generative tissue, limited sperm production is permanent (PSL) and probably has an influence on the reproductive strategies in this species. As nearly no data are available on the life history of H. troglodytes, and in particular information on the phenology of males is lacking, implications of the evolution of PSL in this species are unclear. Nonetheless, our data on other representatives of Austrochilidae (Austrochilus forsteri, Thaida chepu) and Gradungulidae (Progradungula otwayensis) suggest that PSL evolved within Austrochiliodea only in H. troglodytes and might be an adaptation to its troglophilic lifestyle.