Spermiogenesis in three species of cicadas (Hemiptera: Cicadidae)

Spermiogenesis in three species of cicadas representing one cicadettine (Monomatapa matoposa Boulard) and two cicadines (Diceroprocta biconica [Walker] and Kongota punctigera [Walker]) was investigated by light and electron microscopy. Although spermiogenesis was occurring in the testis of adult males of all species, earlier spermiogenic stages were observed in D. biconica only. While spermiogenesis was similar to that described for other insects, some differences were noted. For example granular material did not assemble around the centriole to form a centriolar adjunct but did accumulate in the cytoplasm of early spermatids adjacent to a region of the nuclear membrane where nuclear pores were aggregated. In late spermatids this material accumulated anterior to the mitochondrial derivatives in a developing postero‐lateral nuclear groove. While this material has been named the ‘centriolar adjunct’ by previous authors, its formation away from the centriole raises questions about its true identity. Second, during acrosome maturation an ante‐acrosomal region of cytoplasm develops. Although present in later spermatids, this region is lost in spermatozoa. Interspecific variations in chromatin condensation patterns and the number of microtubule layers encircling the spermatid nucleus during spermiogenesis were noted.     

Semicystic spermatogenesis and biflagellate spermatozoon ultrastructure in the Nile electric catfish Malapterurus electricus (Teleostei: Siluriformes: Malapteruridae)

Spermatogenesis and spermatozoon ultrastructure in the Nile electric catfish Malapterurus electricus are described using scanning and transmission electron microscopy. Although the testis organization conforms to the ‘unrestricted’ spermatogonial type, the species has a rare type of spermatogenesis not previously described among catfishes, ‘semicystic’, in which the cyst ruptures before the spermatozoon stage. Spermiogenesis also involves some peculiar features such as condensation of the chromatin in the posterior part of the nucleus to form a compact electron‐dense mass with some irregular electron‐lucent lacunae, while the uppermost part of the nucleus is a loose electron‐lucent area, absence of the nuclear rotation and, as a consequence, the centriolar complex and the initial segment of each flagellum arise directly in a position perpendicular to the basal pole of the nucleus, and occurrence of numerous vesicles in the midpiece. In addition, spermiogenesis includes migration of the diplosome and mitochondria to the basal pole of the nucleus, formation of two moderate nuclear fossae, each of which contains the centriolar complex, development of two independent flagella and elimination of the excess cytoplasm. The mature spermatozoon has a more or less round head with no acrosome or acrosomal vesicle, a long midpiece with numerous mitochondria and vesicles and two long tails or flagella having the classical axoneme structure of 9 + 2 microtubular doublet pattern and with no lateral fins and membranous compartment. These findings suggest that the ultrastructural features of spermiogenesis and spermatozoa of M. electricus are synapomorphies of types I and II spermiogenesis and spermiogenesis is closely similar to the type described in the Nile catfish Chrysichthys auratus.     

Spermatozoa and spermiogenesis of Holocnemus pluchei (Scopoli, 1763) (Pholcidae, Araneae)

Until now, the knowledge on pholcid spermatozoa is based on two species, Pholcus phalangioides and, incompletely, Holocnemus pluchei. To complete this knowledge and to reveal more potential phylogenetic characters, we have investigated sperm ultrastructure and spermiogenesis of H. pluchei. We found that the sperm cells of this species are clearly different from those of P. phalangioides with respect to: (1) the lack of specialization in the cylindrical acrosomal vacuole; (2) a nuclear canal which is located in the periphery and not in the center of the nucleus; (3) a more prominent postcentriolar elongation of the nucleus; (4) the presence of "inner microtubules" in the implantation fossa in early and mid-spermatids; (5) the absence of a helical band of nuclear material; (6) the proximal centriole which is not prolonged; (7) the types of secretion in the seminal fluid (only two types in H. pluchei). Similarities in the spermatozoa of both species concern: (1) a large implantation fossa which contains large amounts of glycogen in mature spermatozoa; (2) absence of a centriolar adjunct; (3) an axonemal basis located in the posterior part of the implantation fossa; (4) the formation of the so-called cleistospermia in the vas deferens. Our results strongly support systematic relationships within Pholcidae placing these two species in different subgroups.     

Spermatological characters of monozoic tapeworms (Cestoda: Caryophyllidea), including first data on a species from the Indomalayan catfish

The ultrastructure of spermiogenesis and mature spermatozoon in Lytocestus indicus (Cestoda: Lytocestidae) is described; this is the first representative of this group of monozoic, presumably most basal, tapeworms (Eucestoda) from the Indomalayan region to be documented in this manner. Similarly, as in other caryophyllideans, its spermiogenesis involves the formation of a conical differentiation zone with 2 centrioles associated with striated roots and an intercentriolar body. In the course of the process, 1 of the centrioles develops a free flagellum, which fuses with a cytoplasmic protrusion, whereas the other remains oriented in a cytoplasmic bud. Spermiogenesis is also characterized by the presence of electron-dense material in the early stages of spermiogenesis and a slight rotation of the flagellar bud. The mature spermatozoon of L. indicus is a filiform cell tapered at both extremities that lacks mitochondria; its nucleus has parallel disposition to the axoneme and does not reach up to the posterior extremity of the spermatozoon, which is typical for spermatozoa of the type III pattern. The new data confirm that caryophyllideans share the same type of spermiogenesis that is considered to be plesiomorphic in the Eucestoda. The existing information on spermatological ultrastructure of 8 members for 3 of 4 caryophyllidean families from different host groups (cyprinids and catostomids, both Cypriniformes, and mochokids and clariids, both Siluriformes) from 4 zoogeographical regions (Palearctic, Neotropic, Ethiopian, and Indomalayan regions) demonstrates great uniformity in spermiogenesis and sperm ultrastructure, which does not reflect different taxonomic position of the species studied.     

Spermiogenesis and spermatozoon of the tapeworm Ligula intestinalis (Diphyllobothriidea): phylogenetic implications

Using transmission electron microscopy, spermiogenesis and the spermatozoon ultrastructural organization are described in Ligula intestinalis (Linnaeus, 1758) (Diphyllobothriidea), a parasite of the great crested grebe Podiceps cristatus (Linnaeus, 1758). Spermiogenesis starts with the differentiation zone of 2 striated rootlets, 2 centrioles giving rise to 2 flagella, and an intercentriolar body. The latter is composed of 5 electron-dense layers separating 4 electron-lucent layers. In the early stages of spermiogenesis, an electron-dense material is present in the apical region of the differentiation zone. Later, the flagella undergo a rotation and fuse with the cytoplasmic extension in a proximo-distal process. The spermatozoon contains 2 axonemes with a 9 + "1" trepaxonematan pattern, the nucleus, the cortical microtubules, and an electron-dense zone. The spermatozoon anterior extremity in L. intestinalis is characterized by the absence of crested bodies and a ring of electron-dense cortical microtubules. Some characters of spermiogenesis and spermatozoon in L. intestinalis confirm the recent splitting of "Pseudophyllidea" into 2 new orders, i.e., Bothriocephalidea and Diphyllobothriidea. The process of spermiogenesis is similar in both orders for the "type I" of spermiogenesis and the presence of electron-dense material. However, the intercentriolar body is clearly more developed in the Diphyllobothriidea than in the Bothriocephalidea. Moreover, these 2 orders seem to differ in the presence or absence of a ring of electron-dense cortical microtubules in the anterior extremity of the spermatozoon.     

Spermatozoa and spermiogenesis in XyIopIax (Class Concentricycloidea): a new type of spermatozoon in the Echinodermata

In Xylopax turnerae Rowe et al ., 1988 (Class Concentricycloidea, Subphylum Asterozoa) from the Caribbean spermiogenesis culminates in the formation of a complex spermatozoon unlike that of any other echinoderm. Mature testicular spermatozoa are composed of: (1) an elongate, tapered acrosome, segmented internally; (2) a nucleus, finely tapered anteriorly, extremely elongate and rod-shaped posteriorly; (3) a single flagellum attached via a centriolar rootlet to the anterior portion of the nucleus; (4) a single elongate mitochondrion located posterior to the nucleus. Spermatozoa deposited in the ovary have acrosomes which are distinctly granular internally: otherwise these cells resemble testicular spermatozoa. Spermiogenesis differs substantially from the pattern observed in other echinoderms, especially in relation to acrosome development. Examination of alcohol-fixed testes from the New Zealand species Xyloplax medusiforrnis confirmed the presence of filiform spermatozoa, but no details could be established. In contrast to Xyloplax , spermatozoa of Caymanostella sp. (Asteroidea) from a similar habitat to Xyloplax are not modified from those of externally fertilizing Asteroidea, Ophiuroidea, Crinoidea and Holothuroidea. Results lend further support to the existence of a separate class of extant echinoderms (Concentricycloidea) for Xyloplax spp.     

Sperm ultrastructure and a new type of spermiogenesis in two species of Pimelodidae, with a comparative review of sperm ultrastructure in Siluriformes (Teleostei: Ostariophysi)

During spermiogenesis, the spermatids of the pimelodid species Pimelodus maculatus and Pseudoplatystoma fasciatum show a central flagellum development, no rotation of the nucleus, and no nuclear fossa formation, in contrast to all previously described spermatids of Teleostei. These characteristics are interpreted as belonging to a new type of spermiogenesis, named here type III, which is peculiar to the family Pimelodidae. In P. maculatus and P. fasciatum, spermatozoa possess a spherical head and no acrosome; their nucleus contains highly condensed, homogeneous chromatin with small electron-lucent areas; and a nuclear fossa is not present. The centriolar complex lies close to the nucleus. The midpiece is small, has no true cytoplasmic channel, and contains many elongate and interconnected vesicles. Several spherical to oblong mitochondria are located around the centriolar complex. The flagellum displays the classical axoneme (9+2) and no lateral fins. Only minor differences were observed among the pimelodid species and genera. Otherwise, spermiogenesis and spermatozoa in the two species of Pimelodidae studied exhibit many characteristics that are not found in other siluriform families, mainly the type III spermiogenesis.     

Flagellate spermatozoa of Protura (Insecta,Apterygota) are motile

A new model of sperm axoneme with 16 + 0 doublets is described. The spermatozoon of Acerentulus confinis (Apterygota : Protura) has a short conical acrosome, a long helicoidal nucleus, well-developed centriolar adjunct material, and a long flagellum. Using fixation with a glutaraldehyde-tannic acid mixture, without osmium post-fixation, doublet protofilaments, inner dynein arms, radial spokes, nexin bridges, and Y-links of the sperm axoneme of A. confinis and Acerentomon italicum were clearly observed. Optical observation shows that the proturan flagellate spermatozoa are motile cells. The process involving the transformation of the spermatozoa from a coiled to an elongated swimming form was studied by scanning electron microscope. The findings confirmed that flagellar motility is due to the presence of a single dynein arm on doublets in spite of the unusual axonemal pattern.     

Comparative ultrastructure of ant spermatozoa (Formicidae: Hymenoptera)

Mature spermatozoa from spermathecae of founding queens were obtained from 5 species of ants, representing the major subfamilies Myrmicinae (Acromyrmex versicolor, Crematogaster sp.) and Dolichoderinae (Tapinoma sessile, Conomyrma insana, Conomyrma wheeleri). The ultrastructure of ant spermatozoa has many features in common with that of higher insects and is similar to that of other Hymenoptera. Structural similarities to spermatozoa of other Hymenoptera include an acrosome containing an internal rod that extends into the nucleus, two elongate mitochondrial derivatives, a centriolar adjunct, and an axonemal arrangement of 9 + 9 + 2 that includes well-developed coarse, or accessory, tubules. Spermatozoa obtained from A. versicolor, a species that is known to store and utilize viable sperm from this supply for over 10 years, show greater development of the mitochondrial derivatives than do the other species. The most distinctive feature of ant spermatozoa in comparison to other Hymenoptera is the large size of the centriolar adjunct relative to the other organelles. The centriolar adjunct is located posterior to the nucleus, anterior to the mitochondrial derivatives, and opposite the axoneme.     

Spermiogenesis in Psilochorus simoni (Berland, 1911) (Pholcidae, Araneae): evidence for considerable within-family variation in sperm structure and development

A large number of characters and considerable variation among taxa make animal sperm cells promising objects for phylogenetic studies. However, our knowledge about sperm structure and development in spiders is still rudimentary. In pholcids, previous studies of two species representing different subfamily level taxa have revealed conspicuous differences. Here, we report on a representative of a third subfamily level taxon, confirming substantial variation in sperm structure and development within the family. The male genital system in Psilochorus simoni (Berland, 1911) consists of paired testes and deferent ducts which lead into a common ejaculatory duct. The somatic cells of the testes show a high secretory activity, and produce at least two different kinds of secretion. The spermatozoa show features already known from other Pholcidae as well as unique characters. The acrosomal vacuole is tube-like with a narrow subacrosomal space. The axoneme migrates deep into the nucleus and is finally located near the acrosomal vacuole. Thus, the postcentriolar elongation of the nucleus is very long. A centriolar adjunct is not present and after the coiling process the implantation fossa is completely filled with glycogen which is also found in larger amounts within the cytoplasm of the sperm cell. After the coiling process, a vesicular area is present that becomes most prominent in the periphery of the sperm cell and surrounds the axoneme and parts of the nucleus. The secretion sheath surrounding the mature spermatozoon is already formed in the lumen of the testis, possibly by a secretion present in the testis but absent in the deferent duct. Sperm are transferred as cleistospermia. Results are compared with previous studies on pholcid spermiogenesis and sperm structure.     

埃及尼罗河鲶的精子发生和精子的超微结构

用扫描和透射电子显微镜研究了尼罗河鲶——盾头歧须鮠(Synodontis schall)的精子发生和精子的超微结构。精巢中含有无数肾形的生精小叶,我们将其称为"精原无限型"。尽管其精子发生的大体过程与同类鱼无异。但是,在细节上仍具其独特之处。这些特点未见在其他硬骨鱼中报道过。其特点主要是:生精过程中不发生细胞核的旋转,中心粒复合体和轴丝起始段直接发生在核的基底面垂直线上,有无数的粗的固定纤维将近端中心粒和远端中心粒的近侧部连接到细胞核上。另外,精子发生过程中还包括染色质浓缩,细胞质和线粒体向细胞核的尾端迁移,在核的后端中轴位置上形成中等大小的核后凹,近端中心粒和远端中心粒的一部分嵌在核后凹之内,短的胞质内陷管将线粒体与鞭毛分隔开。精子头部接近圆形,无顶体或顶体泡,鞭毛的中段及胞质内陷管均较短,整个鞭毛却很长,鞭毛侧面无翼膜,轴丝呈典型的9 2结构。上述结果显示,盾头歧须鮠的精子发生具有类型Ⅰ和类型Ⅱ的共同派生特征,这种特征在常见的其他硬骨鱼中也是常有的。但是,正如文献所报道过的另两种尼罗河鲶——金鯵(Chrysichthys auratus)和电鲶(Malapterurus electricus)中的情况一样,盾头歧须的精子发生与类型Ⅲ的精子发生过程更为相似。     

Components and development of the centriolar complex during and beyond spermiogenesis in a passeridan bird, the Masked weaver (Ploceus velatus)

The fate of the proximal centriole in passeridan birds is an area of controversy and relative lack of knowledge in avian spermatogenesis and spermatology. This study examines, for the first time, spatiotemporal changes in the centriolar complex in various phases of spermiogenesis in a passerine bird, the Masked weaver (Ploceus velatus). It also describes the configuration of the centriolar complex and the relationship between it and the granular body in both intra- and extra-testicular spermatozoa. It is shown that the proximal centriole is retained and attaches, at its free end, to the granular body of spermatids in every step of spermiogenesis, as well as in mature intra-testicular and post-testicular spermatozoa, including those in the lumen of the seminal glomus. As the centriolar complex, along with its attached granular body, approaches the nucleus in the early spermatid, the proximal centriole articulates with the distal centriole at an acute angle of about 45°, and thereafter, both centrioles, still maintaining this conformation, implant, by means of their articulating proximal ends, at the implantation fossa of the nucleus. In the mature spermatid and spermatozoon, the granular body winds itself helically around the centriolar complex in the neck/midpiece region of the cell, and, thus, becomes the granular helix. The significance of this observation must await future studies, including possible phylogenetic re-evaluation and classification of birds.     

Spermiogenesis and spermatozoon ultrastructure of the cranial digenean Troglotrema acutum (Leuckart, 1842)

Ultrastructure of spermiogenesis and the main characters of the mature spermatozoon of Troglotrema acutum are described by means of transmission electron microscopy. Specimens were obtained from the nasolacrimal sinuses of an American mink (Mustela vison). Spermiogenesis in T. acutum follows the general pattern of digeneans. The zone of differentiation is a conical-shaped area bordered by cortical microtubules and delimited at its base by a ring of arched membranes. This area contains 2 centrioles associated with striated rootlets and an intercentriolar body between them. The centrioles develop 2 free flagella that grow ortogonally to the median cytoplasmic process. The posterior flagellar rotation and proximodistal fusion of the free flagella with the median cytoplasmic process originate the spermatozoon. The mature spermatozoon of T. acutum is characterized by the presence of 2 axonemes of different lengths presenting the 9+'1' trepaxonematan pattern, 2 bundles of parallel cortical microtubules, 2 mitochondria, a nucleus, and granules of glycogen. These ultrastructural characters are compared with other digenean species previously studied and the importance of different spermatological features is discussed.     

Spermatological characteristics of the Trypanorhyncha inferred from new ultrastructural data on species of Tentaculariidae,Eutetrarhynchidae, and Progrillotiidae

The present study focuses on the ultrastructural characteristics of both spermiogenesis and the spermatozoon in the order Trypanorhyncha. New ultrastructural data are presented for two species of the unexplored superfamily Tentacularioidea, Nybelinia queenslandensis, and Kotorella pronosoma. The present study also provides supplementary data on the superfamily Eutetrarhynchoidea, with the analysis of spermiogenesis and spermatozoon of two progrillotiids, Progrillotia dasyatidis and Pro. pastinacae, and new ultrastructural data concerning spermiogenesis in the eutetrarhynchids Dollfusiella spinulifera and Parachristianella trygonis. Spermiogenesis in trypanorhynchs follows the Bâ and Marchand's type I and the ultrastructural organisation of the mature spermatozoon corresponds to the Levron et al.’s type I. The most remarkable characters concerns the number of electron-dense plates constituting the intercentriolar body during spermiogenesis and in the variability of the arc-like row of thick cortical microtubules present in the anterior areas of the spermatozoon because of its variability according to the species.     

Spermiogenesis and sperm in Mesostoma viaregginum (Plathelminthes, Rhabdocoela): an ultrastructural study to infer sperm orientation

Spermiogenesis in Mesostoma viaregginum begins with the formation of a zone of differentiation containing striated rootlets, two centrioles, and an intercentriolar body in-between. These centrioles generate two parallel free-flagella with the 9+“1” pattern of the Trepaxonemata growing out in opposite directions. Spermatid differentiation is characterised by a 90° latero-ventral rotation of flagella and a subsequent disto-proximal centriolar rotation, with a distal cytoplasmic projection. The former rotation involves the compression of a row of cortical microtubules and allows recognising a flagellar side and an aflagellar side in the late spermatid and in the mature spermatozoon. At the end of the differentiation, centrioles and microtubules lie parallel to the spermatid axis. The disto-proximal centriolar rotation is proposed as a synapomorphy for the Rhabdocoela. The modifications of the intercentriolar body during spermiogenesis and the migration of the nucleus and the centrioles towards the cytoplasmic distal projection are also described. The mature spermatozoon of M. viaregginum is filiform and tapered at both ends and presents many features found in the Rhabdocoela gametes. The nucleus disappears before the flagellar insertion and a density gradient of mitochondria is observed along the sperm axis. The anterior end of the spermatozoon of M. viaregginum is characterised by a tapering capped by a membrane expansion. This study has enabled us to describe precisely the orientation of spermatozoa in the Rhabdocoela in general: the centriolar extremity is proposed as the anterior one for the Rhabdocoela.     

Sperm structure and spermiogenesis in Atelura formicaria Heyden (Zygentoma, Insecta)

The spermatozoon of Atelura formicaria (Zygentoma) shows several features that are typical of insects: an apical acrosome, an elongated dense nucleus, a centriole with expanded centriolar adjunct material, two large mitochondrial derivatives, and two thin accessory bodies located beneath the nucleus. The axoneme exhibits a 9 + 9 + 2 pattern with accessory tubules formed by 16 protofilaments and intertubular material. However, spermatozoa of A. formicaria show some remarkable features. The sperm cell is short for an insect, being only 50 µm in length. The nucleus is characterized by the presence of two lateral grooves which are filled with numerous infoldings of the nuclear envelope. In a cross-section the chromatin has the configuration of the Leonardo da Vinci's 'Vitruvian man'. Each mitochondrial derivative has a peculiar structure with peripheral cristae and four crystalline bodies in its matrix; two of these crystalline bodies are large and have differently orientated cristal planes. At the end of spermiogenesis, sperm bundles are stored in the proximal part of the testes. Secretions from the epithelial wall of this region give rise to large globular structures. These include sperm bundles intermingled with dense granules, forming the so called 'spermatolophids'. These formations descend along the deferent duct and are stored in the expanded seminal vesicle. Atelura spermatozoa do not pair as in some Lepismatidae, nor do they fuse as in Tricholepidion (Lepidotrichidae). Thus, sperm aggregation in Zygentoma is realized according to different modalities and can hardly be considered as a synapomorphic trait of its subtaxa.     

Occurrence of biflagellate spermatozoa in Cetopsidae, Aspredinidae, and Nematogenyidae (Teleostei: Ostariophysi: Siluriformes)

In the present study spermiogenesis was investigated in Cetopsis coecutiens (Cetopsidae), and Bunocephalus amazonicus (Aspredinidae), while spermatozoa ultrastructure was investigated in C. coecutiens, B. amazonicus, and Nematogenys inermis (Nematogenyidae). Aspredinidae and Cetopsidae share a spermatogenesis of the semicystic type, and a particular type of spermiogenesis process not reported in any fish group. In the three species analyzed, spermatozoa are biflagellate with flagella having the classical axoneme formulae (9 + 2). The analysis of thirteen characters showed the presence of eight characters shared by Cetopsidae and Aspredinidae, and six characters shared by Cetopsidae and Nematogenyidae, which may suggest that these three families may be more related than actually hypothesized, comprising a very primitive siluriform lineage originated after Diplomystidae.     

Ultrastructure of spermiogenesis and spermatozoa of Gymnotus cf. anguillaris and Brachyhypopomus cf. pinnicaudatus (Teleostei: Gymnotiformes)

The ultrastructure of spermiogenic stages and spermatozoa of representatives of two gymnotiform families, Gymnotus cf. anguillaris (Gymnotidae) and Brachyhypopomus cf. pinnicaudatus (Hypopomidae) were studied. Spermiogenesis of both species is characterized by lateral development of the flagellum and formation of a nuclear fossa. Some differences were found between these species, such as whether (B. cf. pinnicaudatus) or not (G. cf. anguillaris) nuclear rotation occurs, permanence of the cytoplasmic channel, and type and localization of the nuclear fossa. In the G. cf. anguillaris spermatozoon the nucleus is spherical with highly condensed chromatin. The nuclear fossa is shallow and lateral and is associated with the centriolar complex through stabilizing fibrils. The midpiece is short, with many vesicles, a cytoplasmic channel, and elongate mitochondria. In the B. cf. pinnicaudatus spermatozoon the ovoid nucleus is elongated lateral and posterior to the centriolar complex, and has highly condensed chromatin. The eccentric nuclear fossa is of the moderate type, and contains the entire centriolar complex. The midpiece is long, with numerous vesicles, elongate mitochondria, and no cytoplasmic channel. In both species the flagella are laterally disposed in relation to the nucleus and comprise of the classical 9+2 axoneme. Most of the characteristics found in the spermatozoa of these two species of Gymnotiformes are shared with species of Characiformes, whereas only a few are also found in Siluriformes. This suggests that Gymnotiformes and Characiformes may be more closely related than previously proposed.     

Ultrastructure of the spermatozoon of Parachristianella trygonis Dollfus, 1946 (Trypanorhyncha: Eutetrarhynchidae)

The ultrastructural organization of the mature spermatozoon of the trypanorhynch cestode Parachristianella trygonis is described by transmission electron microscopy. The spermatozoon is a long and filiform cell, tapered at both ends, lacking both mitochondrion and crested bodies. Its cytoplasm contains 2 axonemes of the 9+'1' pattern of the Trepaxonemata longitudinally displaced, parallel cortical microtubules, the nucleus and glycogen in form of both alpha-glycogen rosettes and beta-glycogen particles. The anterior extremity of the spermatozoon is characterized by the presence of an arclike row of up to 10 parallel cortical microtubules that partially surround the first axoneme. The present study emphasizes the ultrastructural similarity between mature spermatozoa of all 4 trypanorhynchs that have been studied to date. Nevertheless, several features, i.e., the characteristics of spermatozoa extremities, the absence of crested bodies, and the possible presence of an arclike layer of cortical microtubules, need a more thorough analysis or confirmation in some of these species.