Interrelationships and evolution of the tapeworms (Platyhelminthes: Cestoda)

Interrelationships of the tapeworms (Platyhelminthes: Cestoda) were examined by use of small (SSU) and large (LSU) subunit ribosomal DNA sequences and morphological characters. Fifty new complete SSU sequences were added to 21 sequences previously determined, and 71 new LSU (D1-D3) sequences were determined for the complementary set of taxa representing each of the major lineages of cestodes as currently understood. New sequences were determined for three amphilinidean taxa, but were removed from both alignments due to their excessively high degree of divergence from other cestode sequences. A morphological character matrix coded for supraspecific taxa was constructed by the modification of matrices from recently published studies. Maximum-parsimony (MP) analyses were performed on the LSU, SSU, LSU+SSU, and morphological data partitions, and minimum-evolution (ME) analyses utilizing a general time reversible model of nucleotide substitution including estimates of among-site rate heterogeneity were performed on the molecular data partitions. Resulting topologies were rooted at the node separating the Gyrocotylidea from the Eucestoda. The LSU data were found to be more informative than the SSU data and were more consistent with inferences from morphology, although nodal support was generally weak for most basal nodes. One class of transitions was found to be saturated for comparisons between the most distantly related taxa (gyrocotylideans vs cyclophyllideans and tetrabothriideans). Differences in the topologies resulting from MP and ME analyses were not statistically significant. Nonstrobilate orders formed the basal lineages of trees resulting from analysis of LSU data and morphology. Difossate orders were basal to tetrafossate orders, the latter of which formed a strongly supported clade. A clade including the orders Cyclophyllidea, Nippotaeniidea, and Tetrabothriidea was supported by all data partitions and methods of analysis. Paraphyly of the orders Pseudophyllidea, Tetraphyllidea, and Trypanorhyncha was consistent among the molecular data partitions. Inferences are made regarding a monozoic (nonsegmented) origin of the Eucestoda as represented by the Caryophyllidea and for the evolution of the strobilate and acetabulate/tetrafossate conditions having evolved in a stepwise pattern.     

Adding resolution to ordinal level relationships of tapeworms (Platyhelminthes: Cestoda) with large fragments of mtDNA

The construction of a stable phylogeny for the Cestoda, indicating the interrelationships of recognised orders and other major lineages, has proceeded iteratively since the group first received attention from phylogenetic systematists. Molecular analyses using nuclear ribosomal RNA gene fragments from the small (ssrDNA) and large (lsrDNA) subunits have been used to test competing evolutionary scenarios based on morphological data but could not arbitrate between some key conflicting hypotheses. To the ribosomal data, we have added a contiguous fragment of mitochondrial (mt) genome data (mtDNA) of partial nad1-trnN-trnP-trnI-trnK-nad3-trnS-trnW-cox1-trnT-rrnL-trnC-partial rrnS, spanning 4034-4447 bp, where new data for this region were generated for 18 species. Bayesian analysis of mtDNA and rDNA as nucleotides, and where appropriate as amino acids, demonstrated that these two classes of genes provide complementary signal across the phylogeny. In all analyses, except when using mt amino acids only, the Gyrocotylidea is sister group to all other Cestoda (Nephroposticophora), and Amphilinidea forms the sister group to the Eucestoda. However, an earliest-diverging position of Amphilinidea is strongly supported in the mt amino acid analysis. Amphilinidea exhibit a unique tRNA arrangement (nad1-trnI-trnL2-trnP-trnK-trnV-trnA-trnN-nad3), whereas Gyrocotylidea shares that of the derived lineages, providing additional evidence of the uniqueness of amphilinid genes and genomes. The addition of mtDNA to the rDNA genes supported the Caryophyllidea as the sister group to (Spathebothriidea+remaining Eucestoda), a hypothesis consistently supported by morphology. This relationship suggests a history of step-wise evolutionary transitions from simple monozoic, unsegmented tapeworms to the more familiar polyzoic, externally segmented (strobilate) forms. All our data partitions recovered Haplobothriidea as the sister group to Diphyllobothriidae. The sister-group relationship between Diphyllidea and Trypanorhyncha, as previously established using rDNA, is not supported by the mt data, although it is supported by the combined mt and rDNA analysis. With regards to the more derived taxa, in all except the mt amino acid analysis, the following topology is supported: (Bothriocephalidea (Litobothriidea (Lecanicephalidea (Rhinebothriidea (Tetraphyllidea, (Acanthobothrium, Proteocephalidea), (Nippotaeniidea, Mesocestoididae, Tetrabothriidea, Cyclophyllidea)))))), where the Tetraphyllidea are paraphyletic. Evidence from the mt data provides strong (nucleotides) to moderate (amino acids) support for Tetraphyllidea forming a group to the inclusion of Proteocephalidea, with the latter consistently forming the sister group to Acanthobothrium. The interrelationships among Nippotaeniidea, Mesocestoididae, Tetrabothriidea and Cyclophyllidea remain ambiguous and require further systematic attention. Mitochondrial and nuclear rDNA data provide conflicting signal for certain parts of the cestode tree. In some cases mt data offer results in line with morphological evidence, such as the interrelationships of the early divergent lineages. Also, Tetraphyllidea, although remaining paraphyletic with the inclusion of the Proteocephalidea, does not include the most derived cestodes; a result which has consistently been obtained with rDNA.     

Early morphogenesis in the platyhelminthes with special reference to egg development and development of cestode larvae

Early development of fertilized eggs is reviewed for the main groups within the phylum Platyhelminthes (Turbellaria, Temnocephaloidea, Monogenea, Digenea, Cestodaria and Cestoda). The Turbellaria show two different patterns of embryological development consequent on whether yolk material is contained within the ovum membrane (endolecithal) or whether yolk is added to the outside of the ovum (ectolecithal). In the parasitic groups, those that are oviparous or ovoviviparous produce shelled eggs of the ectolecithal type and, in addition, there are viviparous forms in which egg development shows variations and the amount of yolk, where present, is reduced. In every case examined, however, cleavage is of the spiral type. Within the Cestoda, clear distinction is made between those groups (orders) of worms which are oviparous as compared to those which are viviparous. It is suggested that oviparity is more primitive (ancestral) than viviparity based on adult morphology and on the evidence of life-cycle studies.The continued, post-oncospheral development of cestodes is described as is the ultrastructure of various representative cercoid larval forms. These include: the procercoid of Pseudophyllidea and Haplobothrioidea; the cercoid of Tetraphyllidea; and, within the Cyclophyllidea, the cercoscolex of the Dilepididae, the cysticercoid of the Hymenolepidae, and the cysticercus of the Taeniidae. During post-oncospheral development, in every case examined the posterior part of the developing larva, the cercomer, can be characterized by a microvillar tegument (at least during early development) whereas the more anterior larval body itself, destined to develop in the definitive host, develops microtriches. Modifications to the cercomer, and its microvilli, in the later stages of post-oncospheral development, occur particularly in the more advanced Cyclophyllidea such as the Taeniidae and in some Hymenolepididae.     

Early intrauterine embryonic development of the bothriocephalidean cestode Clestobothrium crassiceps (Rudolphi, 1819), a parasite of the teleost Merluccius merluccius (L., 1758) (Gadiformes: Merlucciidae)

The early intrauterine embryonic development of the bothriocephalidean cestode Clestobothrium crassiceps (Rudolphi, 1819), a parasite of the teleost Merluccius merluccius (L., 1758), was studied by means of light (LM) and transmission electron microscopy (TEM). Contrary to the generic diagnosis given in the CABI Keys to the cestode parasites of vertebrates, the eggs of C. crassiceps, the type of species of Clestobothrium Lühe, 1899, are operculate and embryonated. Our LM and TEM results provide direct evidence that an operculum is present and that the eggs exhibit various stages of intrauterine embryonic development, and in fact represent a good example of early ovoviviparity. The intrauterine eggs of this species are polylecithal and contain numerous vitellocytes, generally ∼ 30, which are pushed to the periphery and remain close to the eggshell, whereas the dividing zygote and later the early embryo remain in the egg centre. During early intrauterine embryonic development, several cleavage divisions take place, which result in the formation of three types of blastomeres, i.e. macro-, meso- and micromeres. These can be readily differentiated at the TEM level, not only by their size, but also by the ultrastructural characteristics of their nuclei and cytoplasmic organelles. The total number of blastomeres in these early embryos, enclosed within the electron-dense eggshells, can be up to ∼ 20 cells of various sizes and characteristics. Mitotic divisions of early blastomeres were frequently observed at both LM and TEM levels. Simultaneously with the mitotic cleavage divisions leading to blastomere multiplication and their rapid differentiation, there is also a deterioration of some blastomeres, mainly micromeres. A similar degeneration of vitellocytes begins even earlier. Both processes show a progressive degeneration of both vitellocytes and micromeres, and are good examples of apoptosis, a process that provides nutritive substances, including lipids, for the developing embryo.     

Vitellogenesis in Echeneibothrium beauchampi Euzet, 1959 (Cestoda: Tetraphyllidea, Phyllobothriidae) (author's transl)]

The vitellogenesis in Echeneibothrium beauchampi is studied by means of electron microscopy. In the follicles, the vitelline cells undergo three progressive changes characterized by cellular size, membrane system development and nature of stored inclusions. 1. Immature cells have a gonial morphology. 2. Maturing cells display a very well developed membrane system. Their cytoplasm is gradually filled with abundant lipid droplets. At the same time, a small number of homogeneous vesicles occur which contain protein material moderately electron dense. Glycogen (alpha and beta) appears at the end of the vitellogenesis. 3. Mature cells show a conspicuous regression of the membrane system. Cellular organelles, nucleus and protein vesicles are found in the peripheral cytoplasmic layer which still contains numerous free ribosomes. The amorphous remaining cytoplasm is filled with large amounts of aggregated lipid droplets which often surround glycogen areas. 4. The presence of numerous lipid droplets and glycogen in the vitelline cells of E. beauchampi shows evidence that this Tetraphyllidea is close to Pseudophyllidea, but the aspect of protein vesicles (shell-protein material) brings it near the Cyclophyllidea.     

Added resolution among ordinal level relationships of tapeworms (Platyhelminthes: Cestoda) with complete small and large subunit nuclear ribosomal RNA genes

The addition of large subunit ribosomal DNA (lsrDNA) to small subunit ribosomal DNA (ssrDNA) has been shown to add resolution to phylogenies at various taxonomic levels for a diversity of phyla. We added nearly complete lsrDNA (4057-4593bp) sequences to ssrDNA (1940-2228bp) for 26 ingroup and 3 outgroup taxa in an attempt to provide an improved ordinal phylogeny for the Cestoda. Ten lsrDNA and seven ssrDNA sequences were generated from new taxa and 13 existing partial lsrDNA sequences were sequenced to completion. The majority of phylogenetic signal in the combined analysis came from lsrDNA (69.6% of parsimonious informative sites, as opposed to 30.4% obtained from ssrDNA), resulting in almost identical topologies for lsrDNA and lsr+ssrDNA (pairwise symmetric distance=6) in model-based analyses. Topology testing found trees based on partial lsrDNA (domains D1-D3)+ssrDNA and complete lsr+ssrDNA to differ significantly; the addition of lsrDNA domains D4-D12 had a significant effect on topology. Overall nodal support was greatest in the combined analysis and weakest for ssrDNA only. Our molecular phylogenies differed significantly from those based on morphology alone. Acetabulate lineages form a monophyletic group, with the Tetraphyllidea being paraphyletic. Support for the combined data was high for the following topology: (Litobothriidea (Lecanicephalidea (Rhinebothrium/Rhodobothrium (Clistobothrium (Pachybothrium(Acanthobothrium Proteocephalidea) (Mesocestoididae, Nippotaeniidea, Cyclophyllidea, Tetrabothriidea)))))); all genus names refer to tetraphyllidean lineages. Although the interrelationships among the four most derived taxa remain uncertain, overall ambiguity of the acetabulate interrelationships was reduced. The Pseudophyllidea were recovered as polyphyletic, with support for a sister-group relationship between Diphyllobothriidae and Haplobothriidea. The monophyly of the Trypanorhyncha was recovered for the first time based on molecular data. The positions of the Trypanorhyncha, Diphyllidea and "Bothriocephaliidea" in relation to other orders remains ambiguous. Higher congruence was found between trees based on model-based phylogenetic methods than with those constructed under the parsimony criterion. Although some uncertainties remain, the addition of lsrDNA D4-D12 has provided an overall more resolved and better supported cestode phylogeny, which further promotes the utility of complete lsrDNA as phylogenetic marker where ssrDNA alone proves inadequate.     

On the phylogenetic positions of the Caryophyllidea, Pseudophyllidea and Proteocephalidea (Eucestoda) inferred from 18S rRNA

A phylogenetic analysis of tapeworms (Eucestoda) based on complete sequences of the 18S rRNA genes of 43 taxa (including new sequences of 12 species) was carried out, with the emphasis on the groups parasitising teleost fish and reptiles. Spathebothriidea and Trypanorhyncha (the latter group being paraphyletic) appeared as basal groups of the Eucestoda but their position was not stable. The tetrafossate orders (Litobothriidea, Lecanicephalidea, Tetraphyllidea, Proteocephalidea, Nippotaeniidea, Tetrabothriidea and Cyclophyllidea) were well separated from the remaining groups. Results supported polyphyly of the Pseudophyllidea formed by two distinct clades: one with diphyllobothriids (Diphyllobothrium, Schistocephalus, Spirometra and Duthiersia) and another including Abothrium, Probothriocephalus, Eubothrium and Bothriocephalus. The former pseudophyllidean clade formed a separate branch with the Caryophyllidea (Khawia and Hunterella) and Haplobothriidea (Haplobothrium), the latter taxon being closely related to either caryophyllideans or diphyllobothriids in different analyses. Proteocephalideans formed a monophyletic group in all analyses and constituted a clade within the Tetraphyllidea thus rendered paraphyletic. Within the Proteocephalidea, the Acanthotaeniinae (Acanthotaenia from reptiles in Africa) and Gangesiinae (Gangesia and Silurotaenia from silurid fish in the Palearctic Region) were separated from parasites of freshwater fish and mammals. The family Proteocephalidae was found to be paraphyletic due to the placement of a monticelliid species, Monticellia sp., in a clade within the former family. The genus Proteocephalus appeared as an artificial assemblage of unrelated taxa which is congruent with previous molecular analyses.     

Vitellogenesis of basal trematode Aspidogaster limacoides(Aspidogastrea: Aspidogastridae)

The vitellogenesis of the trematode Aspidogaster limacoides (Aspidogastrea: Aspidogastridae), a parasite of cyprinid fishes, is described here using transmission electron microscopy. Four different stages of vitellocytes are differentiated: immature vitellocytes, early maturing vitellocytes, advanced maturing vitellocytes and mature vitellocytes. The process follows the same general pattern already described in other free-living neoophorans and parasitic flatworms (i.e. Trematoda, Monogenea and Cestoda): differentiation into mature vitelline cells involves the development of mitochondria, granular endoplasmic reticulum, Golgi complexes, lipid droplets and shell-globules. Mature vitellocytes of A. limacoides are composed of numerous shell-globule clusters, few lipid droplets and glycogen granules. They differ from those of another aspidogastrean Rugogaster hydrolagi in that they possess numerous globules tightly packed and by the presence of only one type of vitelline material. The interstitial tissue of vitelline follicles of A. limacoides contains a peripheral nucleus and long cytoplasmic projections extending between vitelline cells. Since aspidogastreans are considered as an archaic group of parasitic flatworms and thus have a strategic phylogenetic position, future works needs to pay special attention to the ultrastructural and chemical composition of mature vitellocytes within this basal group of trematodes.     

Australicola pectinatus n. gen. and n. sp. (Cestoda: Pseudophyllidea) from the deep-sea fish Beryx splendens from Tasmania

Australicola pectinatus n. gen., n. sp. (Pseudophyllidea: Triaenophoridae) is proposed to accommodate a new cestode from a deep-sea fish, the splendid alfonsino, Beryx splendens Lowe, 1834 (Beryciformes: Berycidae), from the Pacific coast of Tasmania. The new genus is placed in the Triaenophoridae, because it possesses a ventral uterine pore, marginal genital pore, and follicular vitellarium. Australicola is characterized by possessing a massive strobila with very short and wide, markedly craspedote proglottids; vitelline follicles forming a transverse equatorial band; a very deep and narrow genital atrium; a wide, convoluted vaginal canal; and unoperculate eggs. Australicola most closely resembles Eubothrium Nybelin, 1922 and Probothriocephalus Campbell, 1979 in having an unarmed scolex, an unarmed cirrus, the vagina anterior to the cirrus-sac, and cortical vitellaria. It differs from these 2 genera, in addition to the characteristics listed above, in possessing a dendritic rather than an entire ovary. Australicola pectinatus n. sp. is the third cestode described from B. splendens.     

First study of vitellogenesis of the broad fish tapeworm Diphyllobothrium latum (Cestoda,Diphyllobothriidea), a human parasite with extreme fecundity

In the present study, the process of vitellogenesis of one of the most prolific organisms, the broad tapeworm, Diphyllobothrium latum, the causative agent of human diphyllobothriosis, was studied for the first time using transmission electron microscopy. Cytochemical staining with periodic acid-thiosemicarbazide-silver proteinate for detection of glycogen was applied. Starting from the periphery toward the center of the vitelline follicle four stages of vitellocytes are differentiated: immature vitellocytes, early maturing vitellocytes, advanced maturing and mature vitellocytes. Differentiation into mature vitellocytes involves the formation of shell globule clusters containing shell globules, large amount of saturated lipid droplets and glycogen. A peculiar ultrastructural feature of D. latum vitellogenesis is the presence of lamellar bodies in the cytoplasm of mature vitellocytes. This feature is similar to that present in the closely related caryophyllideans and spathebothriideans. Despite the great similarity observed in the embryonic development of diphylobothriideans, caryophyllideans and spathebothriideans, and the fact that their vitellocytes share a feature not reported from other cestode groups, there are substantial differences in the morphology of vitelline clusters, types, amount and localization of their nutritive reserves.     

Spermatological characters of the pseudophyllidean cestode Bothriocephalus scorpii (Müller, 1776)

Spermiogenesis of Bothriocephalus scorpii (Cestoda, Pseudophyllidea) includes an orthogonal development of two flagella, followed by a flagellar rotation and a proximo-distal fusion with the median cytoplasmic process. The fusion occurs at the level of four attachment zones. The presence of dense material in the apical region of the differentiation zone in the early stage of spermiogenesis appears to be a characteristic feature for the Pseudophyllidea. The mature spermatozoon possesses two axonemes of 9+"1" pattern of the Trepaxonemata, nucleus, cortical microtubules, electron-dense granules and crested body. The anterior part of the gamete exhibits a centriole surrounded by electron-dense tubular structures arranged as incomplete spiral. When the crested body disappears, the electron-dense tubular structures are arranged into a ring encircling the axoneme. The electron-dense tubular structures and their arrangement appear to be a specific feature for the clade "Bothriocephalidea". The organization of the posterior extremity of the gamete with the nucleus is described for the first time in the Pseudophyllidea.     

Orders out of chaos – molecular phylogenetics reveals the complexity of shark and stingray tapeworm relationships

Novel molecular data are presented to resolve the long-standing issue of the non-monophyly of the elasmobranch-hosted tapeworm order Tetraphyllidea relative to the other acetabulate eucestode orders. Bayesian inference analyses of various combinations of full ssrDNA, and full or partial lsrDNA (D1–D3), sequence data, which included 134 species representing 97 genera across the 15 eucestode orders, were conducted. New ssrDNA data were generated for 82 species, partial lsrDNA data for 53 species, and full lsrDNA data for 29 species. The monophyly of each of the elasmobranch-hosted orders Cathetocephalidea, Litobothriidea, Lecanicephalidea and Rhinebothriidea was confirmed, as was the non-monophyly of the Tetraphyllidea. Two relatively stable groups of tetraphyllidean taxa emerged and are hereby designated as new orders. The Onchoproteocephalidea n. ord. is established to recognise the integrated nature of one undescribed and 10 described genera of hook-bearing tetraphyllideans, previously placed in the family Onchobothriidae, with the members of the order Proteocephalidea. The Phyllobothriidea n. ord. is established for a subset of 12 non-hooked genera characterised by scoleces bearing four bothridia each with an anterior accessory sucker; most parasitise sharks and have been assigned to the Phyllobothriidae at one time or another. Tentative ordinal placements are suggested for eight additional genera; placements for the remaining tetraphyllidean genera have not yet emerged. We propose that these 17 genera remain in the “Tetraphyllidea”. Among these, particularly labile across analyses were Anthobothrium, Megalonchos, Carpobothrium, Calliobothrium and Caulobothrium. The unique association of Chimaerocestus with holocephalans, rather than with elasmobranchs, appears to represent a host-switching event. Both of the non-elasmobranch hosted clades of acetabulate cestodes (i.e. Proteocephalidea and Cyclophyllidea and their kin) appear to have had their origins with elasmobranch cestodes. Across analyses, the sister group to the clade of “terrestrial” cestode orders was found to be an elasmobranch-hosted genus, as was the sister to the freshwater fish- and tetrapod-hosted Proteocephalidea. Whilst further data are required to resolve outstanding nomenclatural and phylogenetic issues, the present analyses contribute significantly to an understanding of the evolutionary radiation of the entire Cestoda. Clearly, elasmobranch tapeworms comprise the backbone of cestode phylogeny.     

Spermatological characters in Diphyllobothrium latum (Cestoda, Pseudophyllidea)

Spermiogenesis and the ultrastructural features of the spermatozoon of Diphyllobothrium latum (Cestoda, Pseudophyllidea) are described using transmission electron microscopy. Spermiogenesis is characterized by the development of two flagella of unequal length that grow asynchronously. When the first growing flagellum starts to rotate, the second one develops. Flagellar rotation is thus asymmetric and asynchronic. It is followed by proximodistal fusion with the median cytoplasmic process. Electron-dense material is present in the apical region of the zone of differentiation in the early stages of spermiogenesis. The intercentriolar body consists of seven plates: three are electron-dense. Four attachment zones occur in the median cytoplasmic process. An atypical arrangement of striated roots was occasionally observed. The mature spermatozoon possesses two axonemes of 9 + "1" trepaxonematan pattern, nucleus, cortical microtubules, electron-dense granules, and lacks mitochondria. The ultrastructure of the anterior extremity of the spermatozoon in D. latum clearly differs from that in the bothriocephalid pseudophyllideans, mainly in the absence of a crested body and a ring of electron-dense tubular structures. The spermatological data support the assumption that the order Pseudophyllidea is formed by two unrelated clades, "Bothriocephalidea" and "Diphyllobothriidea."     

台湾海峡鱼类绦虫的分子系统学研究

利用DNA测序技术对台湾海峡部分鱼类绦虫的16S rRNA和18S rRNA基因片段序列进行了分析。使用PAUP4·0b10软件构建的进化树显示,目前关于绦虫二叶目、锥吻目、假叶目、盘头目和四叶目的划分是比较合理的,绦虫进化基本遵循了头节形态从简单到复杂的进化规律。报道了国内首次发现的双叶目绦虫,进化树结果初步支持了巨槽属和棘头属的划分。此外,结果也支持了前孔属绦虫的分类地位。但是,对耳槽属绦虫与阶室属绦虫的形态学划分与分子系统学相矛盾,利用16S rRNA基因对盘头目各种的进化树分析与形态学差异很大,这些问题都需要更多研究来进行深入分析。     

Ultrastructural aspects of the germovitellarium of two prorhynchids (Platyhelminthes,Lecithoepitheliata)

Summary

The female gonad of two fresh-water prorhynchids, Geocentrophora baltica and Prorhynchus stagnalis, has been investigated by means of conventional electron microscopy and cytochemical techniques. Both species have an unpaired germovitellarium located under the gut; accessory cells surround the germovitellarium of G. baltica. The germovitellarium consists of a restricted germinative area where early differentiating oocytes and vitellocytes are randomly associated, and an extensive growth area with follicular organization. Each follicle consists of a single alecithal oocyte surrounded by numerous vitellocytes. The main features of oocyte differentiation are the accumulation of lipid droplets and the appearance of Golgi complexes and small bodies possibly representing secondary lysosomes. Vitellocytes show features typical of secretory cells, including well-developed rough endoplasmic reticulum (RER) and Golgi complexes which are involved in the production of type A and type B inclusions, hi both species, type A inclusions appear first, have a glycoprotein content, do not contain polyphenols, and become localized in the peripheral cytoplasm of mature vitellocytes; they have been interpreted as eggshell forming granules. Type B inclusions are larger, have a proteinaceous content with a different structure in the two species examined, and remain scattered in the cytoplasm of mature vitellocytes; they are considered to be yolk. The finding of eggshell forming granules without polyphenols in prorhynchids contrasts with the condition in most platyhelminths that have a sclerotized eggshell formed through a tanning process of polyphenolic substances. The small bodies in the oocytes and the eggshell granules in the vitellocytes of Lecithoepitheliata differ from those observed in prolecithophorans, which have oocyte and vitellocyte inclusions similar to those of the Rhabdocoela.     

A new tetraphyllidean genus and species, Caulopaterapagei n. g., n. sp. (Tetraphyllidea: Phyllobothriidae), from the grey carpetshark Chiloscyllium punctatum Müller & Henle (Orectolobiformes: Hemiscylliidae)

A new genus and species of tetraphyllidean cestode, Caulopatera pagei n. g., n. sp., is described from the grey carpetshark Chiloscyllium punctatum Müller & Henle in Moreton Bay, Australia. The new genus is placed in the Phyllobothriidae, subfamily Phyllobothriinae. Caulopatera n. g. is distinct from all other phyllobothriine genera in having stalked, circular, non-loculate bothridia that lack an apical sucker, testes that are restricted to the region anterior to the cirrus-sac and circum-medullary vitelline follicles. The new genus most closely resembles Carpobothrium Shipley & Hornell, 1906, with which it shares non-loculate, stalked, unhooked bothridia without an accessory sucker and testes that are entirely anterior to the cirrus-sac, but differs from it in that it lacks a slit-like opening in each bothridium and flaps surrounding the opening. The possession of bothridial stalks is consistent with two cestode orders, the Tetraphyllidea and the Rhinebothriidea. The morphology of the bothridial stalks is consistent with other tetraphyllidean genera, in that Caulopatera possesses triangular bothridial stalks surrounding the back of the bothridia, indicating that it belongs in the Tetraphyllidea senso stricto, rather than in the recently recognised Rhinebothriidea.     

Ultrastructural studies on the reproductive system of Gyrocotylidea and Amphilinidea (Cestoda)

Summary The vitellaria, vitellocyte development and vitelloduct of Gyrocotyle urna are described at the ultrastructural level. Vitellar follicles are surrounded by an extracellular lamina; vitellocytes and the periphery of the follicles are enclosed by a cytoplasmic sheath. Immature vitellocytes are spherical and show a high nucleus-to-plasma ratio. During maturation of vitellocytes their cytoplasmic content increases and numerous dictyosomes, endoplasmic reticulum, egg-shell granules and lipid droplets are formed. Lipid droplets and egg-shell granules fill most of the volume of mature vitellocytes. The vitelloduct is ciliated and shows intraepithelial nuclei and intraluminal folds. No cell borders have been found within the vitelloduct. Vitellogenesis and the vitelloduct morphology of Gyrocotyle are compared with those of other parasitic Plathelminthes.Abbreviations I immature vitellocyte - II maturing vitellocyte - III mature vitellocyte - Az cytoplasmic sheath surrounding the vitellocytes and the follicle - bb basal body - ci cilia of the vitelloduct - ld lipid droplet - mi mitochondrium - nu nucleus - rer rough endoplasmic reticulum - sg egg-shell granulum - ve vesicles     

First evidence of the presence of microtriches in the Gyrocotylidea

The Gyrocotylidea, a small and enigmatic group of intestinal parasites of chimaeras, has been considered to be related either to the Monogenea, or, more frequently, to the most primitive monozoic tapeworms (Cestoda), i.e., the Amphilinidea and Caryophyllidea. The present study, based on transmission electron microscopical observations of a species of Gyrocotyle from the rabbit fish, Chimaera monstrosa, in the North Atlantic, demonstrates for the first time the presence of microtriches as surface structures of gyrocotylideans. Because microtriches are considered to be an autapomorphy of tapeworms (Cestoda), in which they differ from other Neodermata (Monogenea and Trematoda), the present data represent another source of evidence in support of a close relationship between the gyrocotylideans and the tapeworms sensu stricto (Eucestoda). Simple morphology, small size, and shape uniformity of the microtriches of Gyrocotyle sp. may indicate they represent an original (plesiomorphic) form that then evolved in more derived cestode groups into a variety of types present mainly on the scolex. The microtriches of Gyrocotyle sp. resemble those found in caryophyllidean, spathebothriidean, pseudophyllidean, and trypanorhynch cestodes, which are considered to represent the most basal groups of the Eucestoda.     

Glycogen synthase in the rat tapeworm, Hymenolepis diminuta--I. Enzyme activity during development and with crowding.

1. Activity of glycogen synthase (E.C. 2.4.1.11) in Hymenolepis diminuta (Cestoda: Cyclophyllidea) was investigated as a function of development and with crowding. 2. Synthase activity was low in the anterior and posterior ends of the worms and highest in the pregravid proglottids in the mid-portion of the strobila. 3. The enzyme activity increased during development of the cestode at least up to 15 days postinfection, but the increase in activity apparently was not due to conversion of the inactive to the active form. 4. Mature oncospheres also contained glycogen synthase, but the activity was lower than in strobilar tissues. 5. Synthase I activities and the proportion of total activity in the I form were generally higher in worms from high density (100 worm) infections than in those from low density (10 worm) infections.