Microtriches of tetraphyllidean metacestodes from Western Mediterranean striped dolphins (Stenella coeruleoalba)

The tegumental structures of two types of tetraphyllidean plerocercoids and two types of merocercoids (Phyllobothrium delphini and Monorygma grimaldii) from Mediterranean striped dolphins, Stenella coeruleoalba, are described for the first time using scanning electron microscopy. The tegument of all of the specimens was fully covered with microtriches. Four basic types were found: filitriches, blade-like spinitriches of different sizes and shapes, cone-shaped spinitriches (with two parallel small projections of equal length at the apex), and crowned cylindrical spinitriches (with 6-7 small papillae forming a crown at the apex); the two latter types are newly described. The two types of plerocercoids had a similar morphology and distribution of microtriches; in addition, cilium-like projections appeared interspersed among the microtriches on the apical sucker, accessory suckers, and distal bothridial loculus. Merocercoids exhibited a greater variety of tegumental structures, especially on the distal bothridial loculus. Both merocercoid types had regularly spaced papillae or "buttons" on the accessory suckers and the distal bothridial loculus that were composed of a central cilium-like projection surrounded by numerous filitriches. However, crowned cylindrical spinitriches were specific to P. delphini and cone-shaped spinitriches were specific to M. grimaldii. Differences in the morphology and distribution of scolex microtriches of adult cestodes have been considered useful for species identification. A previous molecular study has suggested that P. delphini and M. grimaldii are actually different congeneric species. Our study has shown that significant differences in the morphology and distribution of microtriches occur between these species at the merocercoid stage.     

Orygmatobothrium spp. (Cestoda: Tetraphyllidea) from triakid sharks in Argentina: redescription of Orygmatobothrium schmitti and description of a new species

Two species of Orygmatobothrium were found inhabiting triakid sharks collected from the coast of Buenos Aires Province, Argentina. Orygmatobothrium schmittii from Mustelus schmitti is redescribed, including new information on the microtrich pattern. Orygmatobothrium juani n. sp. from Mustelus fasciatus can be distinguished from all other species in the genus using the following combination of characters: worm length, number of proglottids, number of testes, testes distribution, size of eggs, ornamented egg shell, shape of bothridial cleft at level of the marginal accessory sucker, and the extension of vitelline follicles. Species in Orygmatobothrium share a common microtrich pattern with the distal bothridial surface covered with maisiform microtriches interspersed with filiform microtriches, a proximal bothridial surface covered with trifid microtriches, with a medial projection conspicuously larger than the lateral basal projections interspersed with filiform microtriches, an inner and outer surface of the accessory sucker and glandulomuscular organ covered with short filiform microtriches, the scolex proper and cephalic peduncle surface covered with bladelike microtriches, and the germinative zone and entire strobila covered with scutes formed by densely packed filiform microtriches. This general configuration is basically similar to the microtrich pattern described in species of Orectolobicestus and Paraorygmatobothrium.     

Surface ultrastructure of the elasmobranchia parasitizing <Emphasis Type="Italic">Grillotiella exilis</Emphasis> and <Emphasis Type="Italic">Pseudonybelinia odontacantha</Emphasis> (Trypanorhyncha,Cestoda)

The surface ultrastructure of two monotypic trypanorhynch genera is described based on new material of Grillotiella exilis (Linton, 1909) and type material of Pseudonybelinia odontacantha Dollfus 1966. In G. exilis, spiniform microtriches cover the bothrial surfaces and the anterior part of the pars vaginalis posterior to the bothria. Bifurcate microtriches adorn the bothrial margins, filiform microtriches the scolex peduncle, and capilliform microtriches the posterior scolex end. This microthrix pattern resembles that found in, e.g., Grillotia erinaceus (van Beneden, 1858), with the difference that the anterior part of the pars vaginalis is covered with a collar of multidigitate palmate microtriches. The position of Grillotiella within the Grillotiinae, Lacistorhynchidae is supported based on these data. The bothria and scolex peduncle of P. odontacantha are covered with acerosate and unciniform microtriches on the distal bothrial surface and capilliform microtriches on the scolex peduncle. Short filiform microtriches cover the appendix. The microthrix pattern resembles that of the Tentaculariidae but with unciniform and acerosate microtriches densely covering the entire distal bothrial surface. Tegumental grooves are present on the posterior bothrial margin. They can be distinguished from bothrial pits in otobothrioid trypanorhynchs in having similar unciniform microtriches compared to the other parts of the bothrial surface and in lacking any spiniform microtriches. With the absence of bothrial pits as characteristic for the otobothrioids and its characteristic microthrix pattern, P. odontacantha together with Paranybelinia otobothrioides Dollfus 1966, both belonging to the Paranybeliniidae change their position in the most recent system from the Otobothrioidea into the Tentacularioidea.     

Nomimoscolex semenasae n sp (proteocephalidea: Monticelliidae), a cestode parasite of Diplomystes viedmensis (Pisces: Siluriformes) from the Patagonian region of Argentina

Nomimoscolex semenasae n. sp. is described from the primitive fish Diplomystes viedmensis (Siluriformes) from the Patagonian region of Argentina. The new species is placed in Nomimoscolex because of the cortical position of the vitelline follicles, medullary position of the testes, ovary and uterus, and having a scolex with four uniloculate suckers. N. semenasae differs from all other species in the genus by the following combination of characters: (1) apical organ absent; (2) strobila acraspedote; (3) vagina anterior or posterior to cirrus-sac and lacking a sphincter; (4) testes in one irregular layer and in two fields connected anteriorly; (5) uterine stem cortical in immature proglottides, growing from cortical stem into medullary region in mature proglottides; (6) long uteroduct; and (7) presence of spiniform microtriches on all regions of the scolex, proliferation zone and immature proglottis. This is the first record of a proteocephalidean tapeworm in D. viedmensis and in the family Diplomystidae.     

Comparative morphology of surface ultrastructure of diphyllobothriidean tapeworms (Cestoda: Diphyllobothriidea)

Microtriches on the scolices and adjacent strobila of seven species of diphyllobothriidean cestodes (Bothridium pithonis, Cephalochlamys namaquensis, Dibothriocephalus latus, Duthiersia expansa, D. fimbriata, Ligula intestinalis, and Schistocephalus solidus) from different hosts (frogs, snakes, lizards, birds, and mammals) and biogeographic areas were examined using scanning and transmission electron microscopy. The basic structure of the tegument of the seven species studied does not differ markedly from that found in other cestodes. The main characteristic is the presence of electron‐dense bodies and vesicles in the distal cytoplasm. However, this study has shown differences in the morphology of microtriches even among species of the same family. Two different types of microtriches were found, filitriches and spinitriches, with the latter represented by two forms. Our study reveals that capilliform filitriches are most commonly found in Diphyllobothriidea. They were observed mainly on the strobila and the scolices of all but one studied species; individuals of L. intestinalis bore only coniform spinitriches on their surface. The same type of microtriches was found on the cirrus in D. latus. Gladiate spinitriches covered the scolex in both species of Duthiersia, and gladiate spinitriches interspersed with capilliform filitriches were observed on the anterior part of the strobila in D. fimbriata and the posterior part of the scolex in B. pithonis. Individuals of C. namaquensis were covered only by small acicular filitriches. No obvious pattern in the type and distribution of microtriches was observed among species that belong to different families and parasitize distantly related definitive hosts.     

Microthrix pattern of Pseudogilquinia thomasi (Palm, 2000) (Cestoda: Trypanorhyncha) and a review of surface ultrastructure within the family Lacistorhynchidae Guiart, 1927

The microthrix pattern of larvae of Pseudogilquinia thomasi (Palm, 2000) is described for the first time using scanning electron microscopy. The surface ultrastructure of this species consists of three main forms of microtriches: papilliform filitriches, acicular filitriches and quadridigitate to octadigitate palmate spinitriches. The bothria are covered with palmate spinitriches interspersed on some parts with papilliform filitriches. Palmate spinitriches with papilliform, acicular and capilliform filitriches adorn the pars vaginalis and at the anterior part of the pars bulbosa, there is a transition zone in which the palmate spinitriches are replaced by papilliform to acicular filitriches towards the end of the scolex. ANOVA tests with Duncan's post hoc analyses revealed that there are significant differences in the length of spinitriches and their prongs, whether on the surfaces of the bothria or on the surfaces of the scolex peduncle. Callitetrarhynchus gracilis Pintner, 1931 is the second lacistorhynchid species for which surfaces of the segments were examined and the occurrence of large structures called scutes are reported for the first time from this species. There were also significant differences in the base length as well as height of the scutes in different parts of the strobila. In addition to describing the surface ultrastructures of the two lacistorhynchid species, all the available information on the microtriches within the family Lacistorhynchidae is reviewed herein and generic diagnoses related to microtriches are amended. J. Morphol. 277:394–404, 2016. © 2015 Wiley Periodicals, Inc.     

<Emphasis Type="Italic">Luciaella ivanovae</Emphasis> n. g., n. sp. (Eucestoda: Proteocephalidea: Peltidocotylinae), a parasite of <Emphasis Type="Italic">Ageneiosus inermis</Emphasis> (L.) (Siluriformes: Auchenipteridae) in Argentina

Luciaella ivanovae n. g., n. sp. (Proteocephalidea: Peltidocotylinae) is proposed to accommodate a new cestode from a siluroid fish, the manduvé Ageneiosus inermis (Linnaeus) (syn. A. brevifilis Valenciennes) (Auchenipteridae) from the Colastiné River, a tributary of the Paraná River, in Argentina. The new genus is placed in the Peltidocotylinae because it has a medullary ovary with projections into the dorsal cortex, cortical vitelline follicles and testes, a cortical uterine stem, and uterine branches penetrating the medulla and usually as far as the dorsal cortex. Luciaella can be distinguished from all other peltidocotyline genera by the following combination of characters: a quadrangular scolex with four biloculate suckers, each of which has a cap-peak-like structure situated on the anterior margin; the absence of a metascolex; vitelline follicles arranged in two lateral rows which are crescent-shaped in transverse section; testes distributed in two dorsal fields connected anteriorly and sometimes posteriorly; and a vagina which is always anterior to the cirrus-sac and overlaps the convoluted vas deferens. Filiform microtriches are present on all tegumental surfaces, with short filiform microtriches on the scolex and long filiform microtriches on the proliferation zone and on both immature and mature proglottides. The diagnosis of the Peltidocotylinae Woodland, 1934 given by Rego (1994) is amended. L. ivanovae is the fourth proteocephalidean described from A. inermis and the third recorded in this fish host from Argentina.     

Galaxitaenia toloi n. gen., n. sp. (Eucestoda: Pseudophyllidea) from Galaxias platei (Pisces: Osmeriformes, Galaxiidae), in the Patagonian region of Argentina

Galaxitaenia n. gen. (Eucestoda: Pseudophyllidea) is proposed to accommodate Galaxitaenia toloi n. gen., n. sp. from Galaxias platei, a freshwater fish inhabiting Andean lakes in Argentine Patagonia. Galaxitaenia belongs to the Triaenophoridae because it possesses a marginal genital pore, a ventral uterine pore, and a follicular vitelline gland. The new genus can be distinguished from other triaenophorids by the following combination of characters: a scolex with a prominent rectangular apical disc without hooks, grooves, or indentations; neck present; segments wider than long to quadrangular; testes in 2 lateral fields, often connected anteriorly; cirrus unarmed; an ovary situated posteriorly; a vagina posterior to the cirrus sac; vitelline follicles medullary, in 2 ventral fields forming 2 wings interrupted medially; a uterus saccate to branched; and eggs without operculum. The types, distribution, and density of microtriches were analyzed on the surface of the tegument of scolex, neck, and immature segments. The presence of tumuli were observed on the apical disc of the scolex. Galaxitaenia toloi n. sp. is the first triaenophorid species recorded in a South American freshwater fish and the fourth helminth parasitizing the intestine of G. platei.     

Nomimoscolex pertierrae n. sp. (Eucestoda: Proteocephalidea), a parasite of Pseudoplatystoma corruscans (Siluriformes: Pimelodidae) in Brazil and redescription of N. sudobim Woodland, 1935, a parasite of P. fasciatum

The cestode Nomimoscolex sudobim Woodland, 1935 (Proteocephalidea) is redescribed on the basis of its type-specimens and recently collected material found in the intestine of the 'sudobim' Pseudoplatystoma fasciatum (Siluriformes: Pimelodidae) (type-host), from the Amazon River (near the type-locality). Nomimoscolex pertierrae n. sp. is described from the 'surubi' Pseudoplatystoma corruscans from the Paraná River around Porto Rico. Among fifteen known Nomimoscolex species, only N. sudobim possess a cortical uterus in combination with osmoregulatory canals some distance internal to the proglottis margins, as in N. pertierrae n. sp. N. pertierrae differs from N. sudobim, which exhibits the most similarity with the former species, in the morphology of the scolex, the size and unusual shape of microtriches (slender digitiform) on the internal and marginal regions of the suckers, and in the lower mean number of testes [(114 in N. pertierrae, ranging between 216 (recently collected material) and 240 (type-material) in N. sudobim)]. The material described as N. sudobim by Pavanelli & Rego (1992) from the intestine of P. corruscans is considered to be N. pertierrae. The utility of microtriches as characters for distinguishing between species is demonstrated in the clear differences in the shape of those of N. pertierrae and N. sudobim. The morphology of microtriches warrants more attention in future studies on the taxonomy of proteocephalidean cestodes.     

Redescription of Corallobothrium solidum (Cestoda: Proteocephalidea) and erection of a new genus, Essexiella, for tapeworms from channel catfishes (Ictaluridae)

The proteocephalidean tapeworm, Corallobothrium solidum, type species of the genus, is redescribed on the basis of the examination of its type specimens and extensive material recently collected from Malapterurus electricus (type host). Some morphological characteristics of taxonomic importance are reported for the first time, such as the presence of semispherical (U-shaped) sphincters on the external (outer) margin of the suckers, a vaginal sphincter, a well-developed seminal receptacle, and a unique morphology of the eggs. Corallobothrium solidum differs from the 2 remaining species of the genus, both parasitic in channel catfishes (Ictaluridae), in its scolex shape, morphology of its suckers, presence of longitudinal and transverse grooves on the body surface, dense network of excretory canals in the apical part of the scolex, morphology of the eggs, and uterine development. The non-monophyletic nature of Corallobothrium is further supported by molecular data (partial sequences of the 28S rRNA gene) because C. solidum and the 2 remaining species from ictalurids do not form a monophyletic assemblage. Therefore, Essexiella n. gen. is proposed to accommodate Essexiella fimbriatum new comb. (type and only species; syn. Corallobothrium fimbriatum) from channel catfish. Essexiella n. gen. differs from Corallobothrium, Megathylacoides, and Megathylacus by the absence of a sphincter in the suckers, from Corallotaenia by the shape of the scolex and the number and shape of proglottids, and from Paraproteocephalus by the structure of the uterus. The diagnosis of Corallobothrium, which becomes monotypic and restricted to electric catfishes in Africa, is emended. The remaining species of Corallobothrium, Corallobothrium parafimbriatum, is tentatively transferred to Corallotaenia as Corallotaenia parafimbriata n. comb., based on molecular data, small size of the strobila, and shape of the scolex.     

Two new species of Echinobothrium (Cestoidea: Diphyllidea) from myliobatiform elasmobranchs in the Gulf of California, México

Two new species of diphyllidean cestodes, Echinobothrium mexicanum n. sp., and Echinobothrium fautleyae n. sp., are described, the former from the spiral intestines of the bat rays Myliobatis californicus and Myliobatis longirostris and the latter from the spiral intestine of the cownose ray Rhinoptera steindachneri, all caught in the Gulf of California, México. Echinobothrium mexicanum is most easily distinguished from the majority of the species of the genus by the number of large apical hooks in each dorsoventral group (23) and from the remainder in the number of lateral hooklets in each group on the scolex. Echinobothrium fautleyae differs conspicuously from all other members of the genus in that the first and last lateral hooklets in each group are distinctly larger than the other hooklets in each group. Serial sections of the scoleces of these 2 new species confirm that the attachment structures on the scolex are bothria, rather than bothridia, as has been described in other species in the genus, suggesting that the attachment structures of the scolex should be examined in more detail in the genus in general. Palmate microtriches of various forms cover the proximal bothrial surfaces of both new species and are also found on the dorsal bothrial surfaces of E. fautleyae. The distal bothrial surfaces of E. mexicanum are covered with slender spiniform and short filiform microtriches. The cephalic peduncles of both species possess short filiform microtriches only. Comparison of these 2 new species with the other 3 species in this genus that have been examined with SEM suggests that the various forms and distributions of these microtriches is unique in each of these 2 species. We believe these features will prove to be taxonomically useful.     

Anthobothrium lesteri n. sp. (Cestoda: Tetraphyllidea) in Carcharhinus melanopterus from Heron Island, Australia, with comments on its site, mode of attachment, reproductive strategy and membership of the genus

Anthobothrium lesteri n. sp. in Carcharhinus melanopterus from Heron Island, Australia is described and figured. A combination of characters, namely the number of testes, a weakly laciniate strobila and mature proglottis which is conspicuously longer than wide distinguishes it from other species that appear to be consistent with van Beneden's original concept of Anthobothrium . The scolex and the peduncle are relatively small and frail when compared with the rest of the strobila and with the detached proglottis. A myzorhynchus is absent and the stalked, undivided, simple bothridia have thickened margins. Two circular areas of muscles, which resemble accessory suckers, occur in the centre of each bothridium. There is a distinct cephalic peduncle with spiniform blade-like microtriches followed by craspedote proglottides which have a posterior four-lobed velum. Apolysis is well developed and the oncosphere has a polar filament and a dense covering of spines. Comments are made on the site, the mode of attachment, an aspect of the reproductive strategy of the new species in relation to intestinal eversion in some sharks, and membership of the genus.     

Ultrastructure of the tegument of the cestode Paraechinophallus japonicus (Bothriocephalidea: Echinophallidae), a parasite of the bathypelagic fish Psenopsis anomala

Abstract. The ultrastructure of the tegument in Paraechinophallus japonicus (Bothriocephalidea: Echinophallidae), a cestode parasite of the bathypelagic fish Psenopsis anomala , was studied using scanning and transmission electron microscopy. Paraechinophallus japonicus lacks a true scolex. Four different types of microtriches have been observed on the tegumental surface of P. japonicus. Capilliform (∼2.3-μm long) and blade-like spiniform (∼1.4-μm long) microtriches are intermingled on the surface of the pseudoscolex. Capilliform microtriches are distinct in possessing a short base and a long electron-lucent cap. The strobila is covered with two types of microtriches, namely filiform (∼2.1-μm long) and tusk-shaped microtriches (≤4.5-μm long). Tusk-shaped microtriches are limited to the posterior border of each proglottid and are characterized by a short and narrow base, and a large and wide, sharply pointed, electron-dense cap. Similar tusk-shaped microtriches were previously found in members of the family Echinophallidae and may represent an autapomorphy of echinophallid cestodes, all of them being parasitic in centrolophid fish. A unified terminology of microthrix parts is proposed.     

Ultrastructural studies on Diphyllobothrium ditremum and D. dendriticum (Cestoda, Pseudophyllidea), with emphasis on the scolex tegument and the tegument in the area around the genital atrium.

The scolices of D. ditremum and D. dendriticum are similar in regard to general morphology and ultrastructure of the tegument. At the scolex apex and along the upper bothrial edges sensory endings are numerous. The inner bothrium surface of D. ditremum appears to be plain or smooth, the distal cytoplasm is only about 0.6 mum thick and covered with long and slender microtriches. When D. ditremum is attached to the host gut an adhesive layer of secreted substance exists between the inner bothrium surface and the host intestinal villi. In D. dendriticum, however, no secreted layer exists between the inner bothrium surface and the host intestinal villi and the inner bothrium surface of this species appears to be lobed or lappet formed. The lobes are protrusions of the distal cytoplasm and are covered with long and slender microtriches. Large conglomerations of secretion globules appearing to be enclosed by a double membrane, are observed in the scolices of D. dendriticum and D. ditremum, although more frequent in the latter. Differences existing between the distal cytoplasm of the scolex and the gravid proglottid are described. The papillae around the genital atrium in mature and gravid segments were studied. Each papilla appears as a thickening of the distal cytoplasm, which in this region is 10-12 mum thick, when studied with transmission electron microscope. From the area between the papillae sensory endings appearing in sections to be either single, double or triple are described.     

Redescription of <Emphasis Type="Italic">Cetorhinicola acanthocapax</Emphasis> Beveridge &; Campbell, 1988 (Cestoda: Trypanorhyncha) from the Basking Shark <Emphasis Type="Italic">Cetorhinus maximus</Emphasis> (Gunnerus)

Cetorhinicola acanthocapax Beveridge & Campbell, 1988 is redescribed based on adult specimens collected from a basking shark Cetorhinus maximus (Gunnerus) off the east coast of New Zealand. Both the mature and gravid segments are described for the first time, and the first scanning electron micrographs of the armature and scolex microtriches are provided. Novel features include the opening of the vagina anterior to the cirrus-sac and the extension of the vagina anterior to the cirrus-sac. Pectinate microtriches are present on the anterior quarter of the pars pedunculus scolecis scattered between filiform microtriches; the bothrial margins are covered with tridigitate microtriches, while the adherent surface of the bothria have pectinate microtriches. The additional morphological features described are consistent with a close association between Cetorhinicola Beveridge & Campbell, 1988 and the Eutetrarhynchoidea Guiart, 1927.     

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.     

<Emphasis Type="Italic">Nataliella marcelli</Emphasis> n. g., n. sp. (Cestoda: Trypanorhyncha: Rhinoptericolidae) from Hawaiian fishes

A new trypanorhynch cestode from Hawaiian waters is described based on specimens in the collection of the Meguro Parasitological Museum, Tokyo, and newly collected specimens. Nataliella marcelli n. g., n. sp. is characterised by its elongate, craspedote scolex with four small, ear-shaped bothria, an elongate pars vaginalis and long bulbs. The homeoacanthous homeomorphous metabasal armature is comprised of five or six slender, solid hooks per half spiral row. A distinctive basal armature is present, including a combination of six characteristically shaped macrohooks not previously described for trypanorhynchs. The surface ultrastructure consists of gladiate spiniform microtriches that cover the distal and proximal bothrial surface and filiform microtriches on the scolex peduncle. The adults are not known. Nataliella is assigned to the family Rhinoptericolidae Carvajal & Campbell, 1975 based on its possession of four bothria, prebulbar organs and no chainette or intercalary hooks. The family diagnosis is amended to accommodate the new genus. Rhinoptericola megacantha Carvajal & Campbell, 1975 is considered the most closely related species, a position that has been confirmed by molecular analyses of the SSU and LSU rDNA. Nataliella n. g. is intermediate in morphology between the homeoacanth family Tentaculariidae Poche, 1926 (superfamily Tentacularioidea) and the Rhinoptericolidae and other eutetrarhynchid genera (superfamily Eutetrarhynchoidea), sharing characters of both superfamilies.     

Ultrastructure of the cirrus sac of echinophallid tapeworms (Cestoda, Bothriocephalidea) and the terminology of cirrus hard structures

Transmission (TEM) and scanning (SEM) electron microscope methods were used to study the fine structure of the cirrus, cirrus sac, internal seminal vesicle, ejaculatory duct, prostate glands and cirrus armature of Echinophallus wageneri (Monticelli, 1890) and Paraechinophallus japonicus (Yamaguti, 1934) (Bothriocephallidea: Echinophallidae). The cirrus sac of these species has two unique ultrastructural features: a thick wall with two bands of muscles and prominent, rooted hard structures. Rare traits echinophallids share with diphyllobothriideans are microtriches on the ejaculatory duct and with spathebothriideans, well-developed unicellular prostate glands outside the cirrus sac. Because there is a similarity of cirrus armature and rostellar hooks in having a tegumental localisation and in having a heterogenous structure of the blade and root, a cortex, a central pulp region and a recurved apex, these structures are named “modified hooks” instead of spines. They also have a spiral arrangement; no base plate was observed. True spines, as found in trematodes, are between the surface and basal plasma membrane of the external syncytial layer of the tegument, rest on the basal plasma membrane of the distal epithelial cytoplasm, show a homogeneous electron-dark crystalline appearance and are covered by the surface plasma membrane. Aside from the characteristic hooks on the scolex of various cestodes, we see no evidence that would preclude the development of still other specialised structures, such as these modified hooks, from microtriches. In spite of the absence of studies on the development of modified hooks from the cirrus of echinophallids and/or its consideration as derived from microtriches, we assume that like microtriches, formation of modified hooks is from tegumental bodies and therefore they are derivative structures of the cestode tegument.