On the Procercoid Protonephridial Systems of Three Diphyllobothrium Species (Cestoda, Pseudophyllidea) and Janicki's Cercomer Theory

Because the types of hooks are so similar in the oncosphacra/procercoid ef cestodes and in several groups of monogenclic trcmatodes and because the exterior of a procercoid with the hooks in a cercomer is so suggestive of a monogenetic nematode, the development of the procercoids of three Diphyllobothrium species was studied. The intention was to determine whether or not the procercoid protonephridial system would have a developmental stage when its type is similar to, or identical with that type which characterizes the monogeneans. Such a conformity would greatly support the theory of a common origin of monogeneans and ceslodes. However, it has emerged that no similar developmental stage exists. The ontogeny revealed a thorough metamorphosis from a very simple primary protonephridial system (identical with that of the miracidium larva in digeneans) to a secondary system, which develops into the system of the adult tapeworm. This fact may be interpreted as an argument against the supposed inter-relalionships between monogeneans and cestodes. However, the type of hooks and the procercoid cercomer still indicate common ancestors. An analysis of the miracidium, the oncosphaera and the oncomiracidium (the monogenean larva) with reference to their different developmental stages when hatching, gave rise to my interpretation of the fundamental structure of both the miracidium and the oncosphaera as primitively simple and not reduced, Especially the identical type of protonephridial system indicates. in my view, that digeneans and ceslodes originally had a common larva type. If the ceslodes and the monogeneans have common ancestors, then the procercoid may be interpreted as the ontogenetic recapitulation of a common hook-armed ancestor, here named hexucanthoid. This rhabdocoelan creature with six hooks in the cercomer and adapted to an ectocommensalic/ectoparasitic mode of life, is thought to have given rise to the monogeneans, the gyrocotylideans, the amphilinideans and the cestodes. The monogeneans were found to have two fundamentally different types of marginal hooks, and on this basis ihe existence of two different lines of evolution in Monogenea is indicated.     

Phenotypic 'variant' forms of Trichomonas vaginalis trophozoites from cervical neoplasia patients

With the present study, a culture system for successive life-cycle stages of the tapeworm Schistocephalus solidus was developed and this report documents for the first time, cultivation of the procercoid stage of S. solidus from eggs. Additionally we have transformed procercoids dissected from experimentally infected copepods and cultured procercoids into the early plerocercoid stage in vitro. Observations in the culture suggest that the coracidia can interact with their external environment and need no host specific stimuli, except for the components in the culture medium, for activation and hatching from the embryophore. Increasing the culture medium pH from 7.3 to 8.0 improved escape rates and frequencies of hook contractions, suggesting that the oncosphere may recognize and respond to environmental conditions along the host intestine. Procercoids in the culture did not stop growing indicating that conditions within the copepod may be important to limit growth and to induce transformation to plerocercoids. When procercoids are dissected from copepods and transferred to the culture, the outer tegument layers and cercomer starts to loosen. Comparison of the lectin staining of the loosened outer tegument layers and cercomer in procercoids dissected from copepods confirms that transitions of both, the oncosphere to procercoid and procercoid to plerocercoids, has taken place in the in vitro cultures.     

Ultrastructure of the Flame Bulbs and Protonephridial Capillaries of Artioposthia sp. (Platyhelminthes,Tricladida, Geoplanidae)

The protonephridial terminal complex of Artioposthia is formed by one or two terminal cells, each with a nucleus located in the lateral wall of the flame bulb, and probably two proximal canal cells forming the wall of the protonephridial capillary. The weir is restricted to the proximal parts of the flame bulbs and consists of convoluted slits separated by thick cytoplasmic columns. Cross-striated ciliary rootlets running parallel with and obliquely or transversely to the longitudinal axis of the flame bulbs strengthen the walls of the flame bulbs and, to a lesser degree, that of the capillary. Numerous cristate mitochondria are present in the terminal and proximal canal cells. Cytoplasmic processes extend from the terminal cells into the adjacent tissue, and narrow internal leptotriches extend from the cytoplasm of the terminal cells into the lumen of the flame bulbs. The wall of the capillary contains many interconnected, liquid filled spaces that communicate with the lumen of the capillary, and two septate junctions. Phylogenetic implications of the findings are discussed.     

Ultrastructure of the protonephridial system of regenerating Stenostomum sp. (Plathelminthes,Catenulida)

Summary The ultrastructure of the flame bulbs, protonephridial capillaries and duct of fully developed and regenerating Stenostomum sp. is described. Flame bulbs are formed by a single cell whose nucleus is located basally or laterally to the weir. The weir is formed by a single row of transverse ribs connected by a thin membrane, apparently of extracellular matrix. Internal leptotriches arise from the proximal cytoplasm and extend in a (usually) single row along the weir and into the lumen of the distal cytoplasmic tube. Many or all leptotriches do not fuse with the distal cytoplasm. Two cilia are each anchored in the proximal cytoplasm by a cross-striated vertical and lateral rootlet, the latter bent forward and extending for some distance into one of the two cytoplasmic cords along the weir. Each cord contains the lateral rootlet in its proximal part, as well as many microtubules. The distal cytoplasmic tube contains two (longitudinal) junctions, i.e. lines of contact between cell processes of the same, terminal cell. Occasionally, more than two junctions were seen, apparently due to branches of the terminal cell in contact with each other. Flame bulbs join capillaries lined by several canal cells type I, containing few or no microvilli but lateral flames. Such capillaries join a duct (or ducts?) lined by canal cells type II with many long microvilli. The large protonephridial duct is lined by numerous cells with lateral flames and many long microvilli. In regenerating tissue (10.5 hours after cutting) some flame bulbs were free, i.e. not connected to capillaries, and some capillaries openly communicated with the surrounding intercellular space. In the presence of a single row of ribs in the weir, of internal leptotriches, and of vertical and lateral ciliary rootlets, the flame bulb of Stenostomum sp. resembles that of other Plathelminthes much more closely than hitherto thought. The species differs from non-catenulid plathelminths mainly in the large number of glandular cells lining the large protonephridial ducts, in the transverse orientation of the ribs in the weir and in the presence of only two cilia in the flame.     

Porcine model of neurocysticercosis by intracarotid injection of Taenia solium oncospheres: Dose assessment,infection outcomes and serological responses

BackgroundNeurocysticercosis (NCC) is the infection of the human central nervous system (CNS) by Taenia solium larvae that cause significant neurological morbidity. Studies on NCC pathophysiology, host-parasite interactions or therapeutic agents are limited by the lack of suitable animal models. We have previously reported that carotid injection of activated T. solium oncospheres directs parasites into the CNS and consistently reproduces NCC. This study assessed the minimal dose required to consistently obtain NCC by intracarotid oncosphere injection and compared antigen and antibody response profiles by dose-group.Methods/Principal findingsThree groups of pigs were infected with either 2500 (n = 10), 5000 (n = 11), or 10000 (n = 10) oncospheres. Two pigs died during the study. Necropsy exam at day 150 post-infection (PI) demonstrated viable NCC in 21/29 pigs (72.4%), with higher NCC rates with increasing oncosphere doses (4/9 [44.4%], 9/11 [81.8%] and 8/9 [88.9%] for 2500, 5000, and 10000 oncospheres respectively, P for trend = 0.035). CNS cyst burden was also higher in pigs with increasing doses (P for trend = 0.008). Viable and degenerated muscle cysticerci were also found in all pigs, with degenerated cysticerci more frequent in the 2500 oncosphere dose-group. All pigs were positive for circulating parasite antigens on ELISA (Ag-ELISA) from day 14 PI; circulating antigens markedly increased at day 30 PI and remained high with plateau levels in pigs infected with either 5000 or 10000 oncospheres, but not in pigs infected with 2500 oncospheres. Specific antibodies appeared at day 30 PI and were not different between dose-groups.Conclusion/SignificanceIntracarotid injection of 5000 or more oncospheres produces high NCC rates in pigs with CNS cyst burdens like those usually found in human NCC, making this model appropriate for studies on the pathogenesis of NCC and the effects of antiparasitic treatment.     

The fine structure of the protonephridial system in the land nemertean Pantinonemertes californiensis (Rhynchocoela,Enopla, Hoplonemertini)

Summary The protonephridial terminal organ in the nemertean Pantinonemertes californiensis is composed of two cells that are similar in size and shape and are mirror images of each other. Basally in the organ the two cells combine to form a binucleate cytoplasmic mass. Apically they are intimately joined to form a subcylindrical thin-walled weir apparatus; this part is supported by two opposed cytoplasmic columns running the length of the weir region, one originating from each of the two cells, and by a number of regularly spaced circular bars that arise from the two columns. The ciliary flame consists of 94–114 cilia that originate in the bases of the two cells, and it is surrounded by a palisade of incomplete circlets of long, straight microvilli. The convoluted protonephridial tubule is rich in structures that indicate intensive reabsorption from the primary urine. It is argued that the terminal organs in Pantinonemertes and Geonemertes are fundamentally similar and differ only in the amount of microtubules present in the longitudinal supports.Abbreviations BL basal lamina - CF ciliary flame - CT connective tissue - CV coated vesicle - E endocytotic pit - FM filtration membrane - G Golgi complex - LC longitudinal cytoplasmic column - M mitochondrion - MT microtubules - MV microvilli - N nucleus - NPC nucleus of protonephridial capillary cell - PC protonephridial capillary cell - R rootlets - TB transverse bar - TC terminal cell - WE weir, exterior of fenestrated wall - WI weir, interior of same     

Ultrastructural observations of the protonephridia of Polymerurus nodicaudus (Gastrotricha: Paucitubulatina)

Kieneke, A. and Hochberg, R. 2012. Ultrastructural observations of the protonephridia of Polymerurus nodicaudus (Gastrotricha: Paucitubulatina). —Acta Zoologica (Stockholm) 93 : 115–124. We studied different regions of the protonephridia of the limnic gastrotrich Polymerurus nodicaudus by means of light and electron microscopy to determine how freshwater species might differ from their marine relatives. Microscopic and ultrastructural characters are in accordance with another limnic species of Paucitubulatina, Chaetonotus maximus, whose protonephridial system has been previously reconstructed. Shared protonephridial characters of both species include the presence of highly elongate terminal organ cilia, microvilli, and the canal cell lumen as well as the presence of a conspicuous anterior loop of the protonephridial lumen. These features are not present in representatives of earlier, marine, paucitubulatan lineages (i.e., Xenotrichulidae) and so are assessed as evolutionary novelties that were likely important for the successful colonization of the freshwater environment.     

The evolution of the protonephridial terminal organ across Rotifera with particular emphasis on Dicranophorus forcipatus,Encentrum mucronatum and Erignatha clastopis (Rotifera: Dicranophoridae)

Riemann, O. and Ahlrichs, W.H. 2009. The evolution of the protonephridial terminal organ across Rotifera with particular emphasis on Dicranophorus forcipatus, Encentrum mucronatum and Erignatha clastopis (Rotifera: Dicranophoridae). —Acta Zoologica (Stockholm) 91 : 199–211 We report on the ultrastructure of the protonephridial terminal organ in three species of dicranophorid rotifers (Dicranophorus forcipatus, Encentrum mucronatum and Erignatha clastopis). Differences between the three species relate to shape and size, the morphology of the filter region and the number of microvilli and cilia inside the terminal organ. A comparison across Rotifera indicates that the terminal organs in D. forcipatus display a number of plesiomorphic characters, but are modified in E. mucronatum and Er. clastopis. This is in accordance with the results of phylogenetic analyses suggesting a basal position of D. forcipatus compared with the more derived species E. mucronatum and Er. clastopis. Moreover, we survey available data on the terminal organ in Rotifera and discuss its evolutionary transformations. The protonephridial terminal organ in the common ancestor of Rotifera consisted of a cytoplasmic cylinder with cilia united into a vibratile flame and a single circle of circumciliary microvilli. Depending on the topology on which characters are optimized, the site of ultrafiltration was formed by longitudinal cytoplasmic columns spanned by a fine filter diaphragm or by pores in the wall of the terminal organ. In several taxa of Rotifera, the terminal organ – probably independently – lost its circumciliary microvilli.     

Inter- and intra-specific characterization of tapeworms of the genus Diphyllobothrium (Cestoda: Diphyllobothriidea) from Switzerland,using nuclear and mitochondrial DNA targets

Human diphyllobothriosis is caused by at least 14 species of cestodes belonging to the genus Diphyllobothrium. Molecular analysis by sequencing of nuclear and mitochondrial targets identifies some species at inter- and intra-specific level, and helps to reconstruct their phylogenetic relationships. Nevertheless, the suitability of further molecular targets deserves to be widened, and the comparison of samples of different geographical origin could allow their intra-specific characterization, which could also be useful for epidemiological purposes. In this study, we investigated inter- and intra-specific variability among tapeworms of the genus Diphyllobothrium, with focus on Diphyllobothrium latum, originated from Switzerland. Samples were analyzed by comparing the sequences of two nuclear and two mitochondrial DNA targets. We analyzed 27 samples belonging to 4 species (D. latum, Diphyllobothrium nihonkaiense, Diphyllobothrium dendriticum and Diphyllobothrium ditremum), 15 of which isolated from clinical cases (adults and eggs), 2 from wild canines, and 2 from fish of Swiss lakes (plerocercoid larvae); 8 samples of homologous species from other geographic origins were also sequenced and compared with the Swiss ones. Sequences of partial small subunit ribosomal RNA (18S rRNA) gene and partial internal transcribed spacers 1 and 2 (ITS1-2) were not useful even in inter-specific identification, whereas sequences of complete cytochrome c oxidase subunit 1 (cox1) and cytochrome b (cob) genes allowed us to assess inter- and intra-specific variations among the samples. Cox1 and cob could differentiate 3 and 5 haplotypes within the species D. latum. The results are discussed in the light of the anamneses provided by part of the patients.     

Ultrastructure of flame cells and protonephridial capillaries of Craspedella and Didymorchis (Plathelminthes,Rhabdocoela)

Summary The ultrastructure of the flame cells and protonephridial capillaries of the Rhabdocoela Craspedella sp. and Didymorchis sp., ectocommensals on the freshwater crayfish Cherax destructor in eastern Australia is described. The flame cells of both species have variable numbers of cilia without distinct rootlets and with decreasing numbers of axonemal tubules towards the ciliary tips. Bundles of microtubules extend from the cytoplasm adjacent to the ciliary rootlets through the ribs of the weir apparatus into the distal cytoplasmic tube, where the numbers of microtubules gradually decrease. The weir apparatus is formed by a single row of longitudinal ribs connected by a membrane. In Craspedella, but not in Didymorchis, the ribs have external branched leptotriches. Mitochondria are common in the wall of the flame cell of both species. The protonephridial capillary just above the end of the ciliary tuft narrows in both species and bends sharply in Craspedella. The lumen of the flame cell and the capillary is lined by a dark layer of cytoplasm; there is no enlargement of the surface area by microvilli or lamellae. Centrioles were seen in the capillary wall of Craspedella, and in Didymorchis the cytoplasm around the capillaries has a very loose and light appearance. The ultrastructure of the flame cells and capillaries of both species corresponds closely to that of Temnocephala sp.Abbreviations in the figures BB basal body - CE centriole - L leptotrich - M microtubules - ME membrane of weir apparatus - MI mitochondrion - PC protonephridial capillary - R rib (rod) of weir apparatus     

Variation in the reproductive potential of Schistocephalus infected male sticklebacks is associated with 11-ketotestosterone titre

Parasites can impact host reproduction by interfering with host endocrine systems, but the adaptive nature of such effects is disputed. Schistocephalus solidus plerocercoids are parasites of three-spined sticklebacks Gasterosteus aculeatus that are often associated with impaired host reproduction. Here, we relate reproductive behavior and physiology to levels of the androgen 11-ketotestosterone (11KT) in naturally infected and non-infected male sticklebacks from two UK populations. In one population infected males harbored heavy infections and showed uniformly reduced 11KT titres and kidney spiggin (nesting glue protein) content compared to non-infected fish. However in a second population infection levels were more variable and males with smaller infections recorded 11KT and spiggin titres that overlapped those of non-infected fish; among infected males from this population 11KT and kidney spiggin also both correlated negatively with infection severity. Male reproductive behavior correlated closely with 11KT titre in both populations, and infected males with high 11KT levels exhibited normal reproductive behavior. Our results suggest that Schistocephalus infection per se does not block reproductive development in male sticklebacks, and that some male fish may have the ability to breed whilst infected. Our results are not consistent with the hypothesis that Schistocephalus adaptively castrates male hosts via endocrine disruption; rather they support the hypothesis that reproductive disruption is a side effect of the energetic costs of infection.     

Two Human Cases of Diphyllobothrium nihonkaiense Infection in Korea

Diphyllobothrium latum and Diphyllobothrium nihonkaiense are the 2 reported main causes of human diphyllobothriasis in the Republic of Korea. However, the differentiation of these 2 species based on morphologic features alone is difficult. The authors used nucleotide sequencing of the mitochondrial cytochrome c oxidase subunit 1 (cox1) gene to diagnose Diphyllobothrium spp. Two patients visited the emergency room at Kyungpook National University Hospital on 3 April and 12 April 2013, respectively, with fragments of parasites found while defecating. The parasites were identified as Diphyllobothrium spp. based on morphologic characteristics, and subsequent cox1 gene sequencing showed 99.9% similarity (1,478/1,480 bp) with D. nihonkaiense. Our findings support the hypothesis that D. nihonkaiense is a dominant species in Korea.     

Ultrastructural reconstruction of Taenia ovis oncospheres from serial sections

The cellular organisation of Taenia ovis oncospheres is interpreted from ultrathin serial sections and transmission electron microscopy following high pressure freezing and freeze-substitution. The surface of a hatched, non-activated T. ovis oncosphere is covered by an oncospheral membrane below which is the tegument bearing microvilli. The basal lamina of the tegument is underlain by broad bands of peripheral somatic musculature. Three pairs of hooks and associated muscles are present in the somatophoric third of the oncosphere. Approximately 19 cells of seven different types were identified which include: (i) a quadri-nucleated syncytium of penetration gland type 1 containing two lateral pairs of cell bodies interconnected by narrow cytoplasmic bridges (PG1); (ii) a quadri-nucleated syncytium of penetration gland type 2 (PG2); (iii) a single-nucleated median mesophoric gland cell; (iv) 10 somatic cells; (v) two germinative cells; (vi) two nerve cells; and (vii) a pair of median somatophoric cells. This study provides a clear understanding of the morphology of T. ovis oncospheres and forms the basis for further investigations into the biology of taeniid oncospheres.     

Diphyllobothriids (Cestoda: Pseudophyllidea) from the long-finned pilot whale Globicephala melas (Traill, 1809) off the Faroe Islands,with comments on the taxonomy of Diphyllobothrium Cobbold, 1858 species from cetaceans

The taxonomy of marine species of the genus Diphyllobothrium, particularly those parasitic in cetaceans, is rather confused. During parasitological investigations of long-finned pilot whales Globicephala melas from waters off the Faroe Islands, five diphyllobothriid species were detected: Diphyllobothrium sp. (possibly D. polyrugosum), D. stemmacephalum, Diphyllobothrium sp. A, Diphyllobothrium sp. B and Diphyllobothriidae sp. D. stemmacephalum is reported for the first time from G. melas. The stituation regarding the taxonomy of Diphylobothrium species from cetaceans is briefly reviewed. It is concluded that the recent development of genetic techniques may be of great value in relation to clarifying the taxonomy of this group.     

Development of the excretory system in the polyplacophoran mollusc,Lepidochitona corrugata: The protonephridium

A single pair of protonephridia is the typical larval excretory organ of molluscs. Their presence in postlarval developmental stages was discovered only recently. We found that the protonephridia of the polyplacophoran mollusc, Lepidochitona corrugata, achieve their most elaborate differentiation and become largest during the postlarval period. This study describes the protonephridia of L. corrugata using light and electron microscopy and interactive three‐dimensional visualization. We focus on the postlarval developmental period, in which the protonephridia consist of three parts: the terminal part with the ultrafiltration sites at the distal end, the voluminous protonephridial kidney, and the efferent nephroduct leading to the nephropore. The ultrafiltration sites show filtration slits between regularly arranged thin pedicles. The ciliary flame originates from both the terminal cell and the duct cells of the terminal portion. The efferent duct also shows ciliation. The most conspicuous structures, the protonephridial kidneys, are voluminous swellings composed of reabsorptive cells (“nephrocytes”). These cells exhibit strong vacuolization and an infolding system increasing the basal surface. The protonephridial kidneys, previously not reported at such a level of organization in molluscs, strikingly resemble (metanephridial) kidneys of adult molluscan excretory systems. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Localisation of three host-protective oncospheral antigens of Taenia ovis

Immunohistochemistry, confocal immunofluorescence and immunogold labelling were used to determine the localisation of the host-protective antigens To16, To18 and To45W in Taenia ovis oncospheres. During maturation of the adult tapeworm the antigens were initially seen as diffuse staining in the developing oncospheres but in mature oncospheres four distinct cells stained positively for the antigens. Confocal fluorescence microscopy using different fluorophores revealed that each of the antigens co-localises within the same cells in the oncosphere. No surface localisation was seen in non-activated or recently activated parasites. Immunogold labelling of non-activated oncosphere sections viewed in transmission electron microscopy revealed labelling of bilateral cells, however the identities of these cells was unclear due to deficiencies in the current level of understanding of oncosphere ultrastructure. Localisation of all the antigens changed dramatically after oncospheres were activated in vitro with each of the antigens being dispersed more generally throughout the parasite parenchyma. During development of the parasites in in vitro culture, surface localisation of the proteins was seen in parasites after 3 or more days in culture. All three antigens were found to be completely absent in parasites by 15 days of culture. The location of the host-protective antigens suggests that initially the invading oncospheres are not susceptible to vaccine-induced antibody and complement mediated attack, but that as the parasites mature, the host-protective antigens come to be associated with the parasite’s surface, rendering them susceptible to immune attack.     

Ultrastructure of the protonephridial system of Polystomoides malayi Rohde and P. renschi Rohde (Monogenea: Polystomatidae)

Rohde K. 1973. Ultrastructure of the protonephridial system of Polystomoides malayi Rohde and P. renschi Rohde (Monogenea : Polystomatidae). International Journal for Parasitology3: 329–333. Polystomoides malayi and P. renschi have three types of protonephridial flames. The first type is a typical flame cell with internal and external ribs connected by a weir membrane without nephrostomes, and with internal and external leptotriches. The second type is a flame cell complex consisting of at least two flames reaching into a common cavity. The third type is a non-terminal (= lateral) flame in the protonephridial ducts, consisting of loosely arranged cilia many of which have lateral tube-like extensions and whose tips have irregularly arranged filaments gradually decreasing in number. The number of cilia in all types of flames varies. The smallest capillaries are strongly convoluted and have a smooth or slightly reticulated surface, the larger ducts have strongly reticulated walls and single cilia may be found in the cavities of the reticulum.     

Echinococcus granulosus: use of an intermediate host mouse model to evaluate sources of protective antigens and a role for antibody in the immune response.

A Balb/cJ mouse model was used to determine which stage of the E. granulosus life cycle possessed the most potent protective antigens. Mice were immunized with crude extracts of protoscoleces, brood capsules, cyst fluid, adult worm tissue, eggs or oncospheres and then challenged intraperitoneally with 600 activated oncospheres. Sonically disrupted oncospheres induced the highest levels of protection (greater than 90%) at doses greater than or equal to 10(3) oncosphere equivalents per mouse. High levels of protection were maintained when these preparations were solubilized in SDS. Immunization with Taenia ovis or T. hydatigena oncosphere preparations induced a maximum of 62 and 40% cross-protection, respectively. In passive transfer experiments, serum from triple-infected immune donors that were completely resistant to subsequent challenge induced 69% protection in naive recipients (P less than 0.01). Serum from mice that had been immunized with oncosphere sonicates that were shown to be highly immune, failed to induce statistically significant protection in recipients. A sheep trial confirmed the protective ability of prior infections. Immunization of sheep with a SDS solubilized oncosphere preparation produced 91% protection (P less than 0.01).     

A review of protonephridial excretory systems in Acanthocephala

Our present understanding of the excretory system of Acanthocephala is largely the result of work done by 5 German scholars: Kaiser, Schepotieff, Meyer, Kilian, and von Haffner. Present studies indicate that a protonephridial system is restricted to the family Oligacanthorhynchidae. However, many members of this family have not had a protonephridial system described. Three nephridial designs have been described: 1) dendritic type, organized as branches of a tree where each final branch terminates in a ciliated bulb; 2) capsular type, in which all ciliated bulbs empty directly into a common chamber; and 3) rudimentary type, consisting of a single cell with a patent ciliary pouch but no ducts to the outside. The first 2 types are a syncytia with 3 nuclei located in the capsule or stem wall and none in the flame bulbs. These excretory systems consist of 2 clusters of flame bulbs that empty separately into an expandable excretory bladder which in turn empties into ducts of the reproductive system. This urogenital system empties to the outside through a gonopore located at the tip of the penis in males and the posterior terminus of the vagina in females. Cilia occur in certain excretory tubes, depending on sex and species, but are unknown in the excretory bladder or ducts leading into it. The rudimentary type consists of a cell whose posterior extension terminates near the bursal lumen, but it is not known if this is significant for the discharge of material. There is no information on the physiology or biochemistry of the excretory system or its products.